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2 '! r,, H,, Ji ' i! :t: ;! :j.:ti vity (f m 00,~ _.\,,!i :i! 953. Pro~rc:-' :n! :.:cmra Aaska Tr:tn )52. H. Moose and tire ' Eco. Conf. 3: -3:: p3. Wof report. Fed ep. W-7-4. Aask.~ :>p. :)tin ek-shiras m 00,_ J6. -/.Jose and deer winter fj. anage. 3: ~ by moose and white! i :jcervid and snowshoe.,;i 38: n!ft9ry methods: a re.foeha) ior of moose in. G::unt! Bio. 3:03-., ~~97!. Radioteemetry f\innesota. J. Wid!.,.jf:' of.~aptive Michigan ~ 'iga of Aaska. Quar- ~~~ A Effects of : a ~erature review.,3ureau of Land ~an- oose coactions with. 56. : se Royae moose od browse by moose r Aaska. U.S.D.A.,!g. 7pp., competition during.. Moose Conf. and t of Fish and Game ' versity Press 47 Caribou Rangifer tarandus NOMENCLATURE COMMON NAMES. CaribmJ, reindeer, "deer" SCENTFC NAME. Rangifer tarandus SUBSPECES. R. t. groenandicus, barren-ground caribou or American tundra reindeer: R. t. granti, Aaskan barren-ground caribou or Grant's caribou: R. r. caribou, Amerh:an woodand caribou or woodand caribou; R. t. pearyi, Peary caribou or Peary reindeer (Banfied 96). The Queen Charotte sand's caribou (R.. dawsoni) probaby became extinct shorty after 90 (Banfied 963). be cau5t:s for its extinction are unknown, but it is s.uggested that habitat deterioration through ameioration of L"te cimate and oss of genetic pasticity through isoation were more important than hunting or other human interference (Banfied 963). The caribou is the ony North American cervid that has estabished year-round popuations north of t~e rreeine. into some of the harshest ands in North America. The caribou frrst appeared in Europe in the mid-peistocene about years ago and is a primitive member of the deer famiy Cervidae (Banfied ). n North America. caribou were probaby in Aaska and the Yukon before the Wisconsin gaciation, but their earier occurrences and origin are, as yet, undescribed (Banfied 96). The features that suggest the primitive nature of Rangifer incude the possession of anters by both sexes, ong metapodia bones, we-marked tarsa and interdigita gands, and reativey simpe crests on the cheek teeth (Banfied 974). Banfied's (96) revision of Rangifer is an exc~cnt source for the Paeontoogica record. vernacuar names, previous revisions, and the current taxonomy. Suppementary information is in Kesa (968). DSTRBUTON The Peary caribou has ranged as far south as the mainand coast of t' Arctic Ocean, and on occasion has intergraded me Canadian mainand form of barren-ground caribou (Banfied 954b, 96; Manning 960). n tum, some mainand barren-ground ~~---~ "' Frank L. Mier FGURE Distributmr. of caribou ( Rang{fer rarandus spp.):, R. r. caribou: 2. R. r. daw.wmi, E =extinct; 3, R. r. grami: 4. R.. groen/andicus; 5. R.. pearyi. The actua boundaries between the \Ubspecies are not we defined: seasona overap did and probaby sti does occur in many area!>. Arrows indicate areas of probabe subspecific overap. After Banfied 974. caribou have migrated north to the more southery isands of the arctic archipeago and shared portions of summer ranges of Peary caribou. Most of the interchange had apparenty ceased by the end of the 920s, a period during which native hunters took excessive harvests from migrating herds (Hoare 927: Manning 960). Litte is known of the persistence of movements between southern isands and the coasta mainand and of the present genetic interchange between Peary and barren-ground caribou. On the Canadian arctic archipeago, the Peary caribou shares its range with ony one other unguate, the muskox (Ovibos moschatus) (Tener 963). The tota number of Peary caribou in the Canadian High Arctic probaby does not exceed 0, (Gunn et a. 98). Currenty, Mevie and Prince Patrick isands-the Queen Eizabeth group-are the heartand of Peary caribi... north of 70 0<)'' N atitude 923 f,_

3 924 ARTODACTYLA (Mier et a. a). Peary caribou on the western Queen Eizabeth hands decined from about in 96 (Tener 963 i by about 89 percent in 974 (Mier et a. a). Peary caribou supposediy occur in higher numbers on Banks, Victoria. Prince of Waes. and Somerset isands (Mier 978) (figure 47.). The barren-ground caribou of Aaska and Canada ranges over thousands of square kiometers of arctic tundra. Some herds remain on the tundra year-round, whie others migrate south into the borea forest (taiga} for the winter. Those barren-ground caribou that are m the boreai forests in autumn may intergrade with northem popuations of woodand caribou. Barren-ground carihou share portions of their range with moose (A ees aces) and muskoxen. n Canada the barren-ground caribou occurs on the arctic isands of Baffin. Byot, Southampton, and Coa 3, and possiby sti on Victoria and King Wiiam {Banfied 96). On the mainand it occurs from west of Hudson Bay in the District of Keewatin, Northwest Territories. and orthern "Manitoba westward :Jew- the District of M:tckenzie. Northwest Territore northern Saskatchewan. and occa~ionay nonheasttt Aberta to the Mackenzie River. Northwest Terrnont> (fig. 47.). The barren-ground caribou of the Northwest T~r ritorie~ occur in nine herds or popuatiom. CaL! 978). Those incude the barren-ground caribou th~t seasonay range into tht! provinces to the south. Recent estimates of densities based on aeria surveys (Cak: 978) suggest that there are curr~nry ahdm 550J) barren-ground caribou west of Hudson Bay and east ot the Mackenzie River (abe 47.). At the Mackenzie River the Canadian form of barren-ground caribou (groenandicus) gives way tc' and sometimes intergrad~s with the Aaskan form ot barren-grou!d caribtjy (granti). T:e Aaskan barrenground caribqij.ranges west and north across the Yukon Territory. onto the north sope of Aaska. and throughout arctic Aaska. ts surviva as a pure stock on the TABLE 47.. Recent estimates of sizes of major herds of caribou ( Rangifer tarandus) in North America Herd Aaska" Adak Aaska Pen~nsua Andreafsky Beaver Centra arctic Chis ana Deta Fonymie Granite Mountain Kenai Kibuck Mountain Macomb McKiney Memasta Muchatna Nechina Rainy Pass/Farewe Ray M0untains Sunshine-Coudy Mountains Teshekpuk Porcupine Western arctic Canada' Parcupin:'' Buenose Bathurst Bevery Kaminuriak Wagner Bay Mevie Peninsua Lorriard Baffin sancf George River Estimae b.500-5, ,000, , , , , ,000 75,000 93,000-00, ()t)(} 50,000 94,000 38,000 29,000 52,000 7,000 20,000 85,000 Year i 9' Trend stabe increasing unknown stabe unknown stabe decreasing decreasing unknown increasing unknown unknown decreasing stabe increasing decreasing unknown unknown unknown unknown stabe decreasing stabe increasing decreasing decreasing decreasing increasing increasing unknown decreasing increasing "Data taken from Davis (978). t> Extrapoated from 975 survey data (Davis 978).,. Porcupine estimates from Aaska and from Canada are for the same herd. d Data taken from Caef (978); except Bevery and Kaminuriak estimates from A. Gunn (persona communication) and George River estimate from S. Luttich (persona communication). Aas! of e) reind barre with divid pace abou sequ< abou don pred and habi: Ne\\ Ne\\ and com tion war virg ike port beer the tem affe Ne Riv Ga~ A t Se 9- Sta; car Lui the. Yu\ Th. Pe: car de ze me ~ ~ " ~- --~-- ~ ~-- D: C ~.~}.: a!-- m_ a ~P:f:~_; c c

4 CARBOU (Rarzgifer tarandus) 925 :west Terrnont i i ~ onhwesr Tt r pations t Cuk \d caribou tha: :e south. Recen. '! surv~ys t C:t!,, ubou~ 550 Ot...-'., ~jbay and east ot ' ' tadian form of -( gives way to Jia5kan form of askan barren-.ross the Yukon. and throughe stock on the, k '. nenca ~~----- Trend stabe increasing 'Unknown" stabe unknown stabe decreasing decreasing unknown increasing unknown unknown decreasing stabe - mcreasng decreasing unknown unknown unknown unknown srabe decreasing stabe increasing decreasing decreasing decreasing increasing increasing unknown decreasing ttcreasing 'micat'i'on) and Aaskan Peninsua is questionabe. however,!:. ecause of extensive interbreeding with introduced domestic reindeer (Banfied 96). The Abskan form of barren-ground caribou (and some ikey intergrades with woodand and/or free-ranging reindeer) has been divided into 2:! herds (Davis 978). Earier estimates paced the number of Aaskan barren-ground caribou at about 600,000 (Skoog 968~ Hemming 97). Subsequent work (tabe 47.) indicated a marked decine to about (Davis 978). Overhunting :md predation by woves (Canis upus) have been suggested as predominent causes for that decine. The woodand caribou occupies the borea forest and apine tundra extensions of suitabe mountainous habitats. Woodand caribou once occurred in Maine, New Hampshire, Vermont, Michigan. Minnesota. New Brunswick, Nova Scotia, and Prince Edward sand. The range ofthe woodand caribou has decreased consideraby since the 800s, probaby due to destruction of the cimax forests and overhunting. The northward extension of white-taied deer ( Odocoieus virginianus) foiowing the oss of the cimax forests ikey eiminated woodand caribou from the southern ponion of their range. An additiona factor may have been the deer's roe as a carrier of moose sickness," the parasitic meningea worm (Pareaphosrrongyus renuis) (Smith et a. 964: Anderson 97) that aso affects woodand caribou. Now their range has been pushed nonhward to Newfoundand, Quebec, nonh of the St. Lawrence River. except in the Shickshock Mountains of the Gaspe Peninsua, Ontario, Manitoba, Saskatchewan, Abena, remnant popuations in th~ Samo River Sekirk area (Freddy and. Erickson 975: Johnson 976) of daho. and possiby Montana, Washington State, and British Coumbia (fig. 47. ). The woodand caribou range extends nonh in the east to the Ungava Labrador Peninsua and Newfoundand. in the west to the District of Mackenzie, Northwest erritories, the Yukon Territory. and the Copper River area of Aaska. The woodand caribou no onger exists on the Kenai Peninsua (Banfied 96). The ranges of woodand caribou are shared with 3evera other members of t~!e deer famiy: white-taied deer, back-taied deer ( 0. hemionus coumbianus), mue deer (0. h. hemionus), moose, and ek (Cervus eaphus). DESCRPTON Caribou are medium-sized deer with reativey ong egs, arge hooves, and broad muzzes, Their heads are eongated, with forehead-nose profies that vary from amost straight to the 'roman noses'' of some mature. maes. The muzze is bunt, except in Peary caribou, and we haired except for the sma ova rhinarium (Pocock 923). The physica appearances of caribou differ among subspecies; many of the differences are given in Banfied (96 ). Detaied descriptions of caribou are given by Dugmore (93) for woodand caribou of Newfoundand, by Murie (935) for Aaskan-Yukon caribou, and by Harper (955) and Kesa (968) for barren-ground caribou of the Northwest Territories. Size. Maes weigh from about 0 kiogram!> kg for Peary caribou (season unknown. Banfied 96 to 299 kg for Aaskan barren-ground caribou prerut. Sko0g 968). Seasona difference~ in body weight:.. especiay for adut maes. are so great that much of the range of differences between maes and femaes i~ ost. Generay. however. mature femae caribou are ahout 0-5 per~em smaer and weigh 0-50 percent e:-.~ ihari uduit mae$. Peage. The forest-dweing woodand caribou have the darkest peage and the Peary caribou the ightest. with more gray than brown in their coats. Barrenground caribou have many variations in intermediate shades of coor reative to the other two subspecies. The underparts of caribou are ighter. eg cooring and socks are pronounced, and fank stripes vary from prominent to acking. Descriptions of peages using standard coor guides are given in Manning ( 960) and Banfied (96). Each spring, caribou ose their winter pertge. which fas away in patches, reveaing darker hair beneath and eaving the caribou with a ragged appearance. Prime bus are usuay the first to deveop seek summer coats. foowed by juvenies and yearings, by cafess cows, and, ast, by materna cows. By,..ugust, a but cows in poor condition are in nt>w coats. which are the darkest. n autumn, the hair engthens and onger, white-tipped guard hairs grow to form the winter coat. By prerut (September-October). the bus have deveoped handsome white manes and the peage shows the maximum contrast of an individua's cooration. As winter progresses, the guard hairs are beached and many of the hair tips are broken off. resuting in a genera ightening and Joss of contrast in the peage. Finay, the winter coat fas away and the cyce begins again. Caves are born with a ight brown or red brown body and a dark brown to back dorsa stripe from the neck to the.tai. Their undersides are white or ight gray (Kesa 968). Anters. Both maes and femaes have anters, athough the mae's are arger and can be of impressive proponions. The main beams are ong and curved toward the anima's posterior with a cyindrica to fat- t..n~.d. cross-section. Usuay one and sometimes two brow tines are widey pama ted (the 'shoves'') and the dominant one extends venicay over the face. The brow and second tines extend anteriory and the termina tines extend posteriory, and can be either distinct or pamate. The anters of femaes and young caribou are smaer a'd simper. Femaes do, however. sometimes grow miniature repicas of the mae's arge anters. Caves grow "spike" anters that remain in vevet for their first winter and spring. The shapes of caribou anters vary greaty, and no two sets are the same. One anter of a pair is not the mirror image of the other (Banfied 954a). The varia-

5 926 ARTODACTYLA ---- tion among popuations usuay negates the use of anters in detaied taxonomic work (Banfied 96 >. athough Bubenik ( 975h argued for their potentia use in taxonomy. The annua cyce of anter deveopment (e.g., Banfied 954a; Harper 955: Moisan 959: Bubenik 975/J varies with the sex. reproductive status. and age of the caribou. Maes shed tht'!ir anters during or after the rut. The roe of testosterone in the anter cyce of caribou is minor. and under det!p physica stress some oder buhc;; can drop their anters before the end of the rut (Murie 935; Bubenik 975b). Those bus without anters immediatey ose their sex drive and dominance and do not further participate in rutting activities (Espmark 964a). The anteress bus turn to foraging and recover their physica condition more rapidy than other prime bus that participated in th~ entire rut; such a process coud have surviva vaue. Femae caribou retain their anters through the winter and often shed them about the time of caving (e.g., Lent 965q; Skoog 968; Bergerud 976). The presence of osteoytic resorption in anters in December suggests that the retained anters of femaes and juvenies coud serve as a cacium bank during the winter (Beanger et a. 967). The annua cyce of anth'!r vevet and anter shedding varies with sex and age. Bergerud (976) iustrated the anter cyce for woodand caribou in Newfoundand. Skoog (968) and Mier (unpubished data) observd sight subspecific variations from woodand caribou in the annua a.nter cyce of barrenground caribou: () mature ban en~ground bus begin to grow anters in eary March, athough a few carry hard anters unti ate February; (2) yearing barrenground caribou begin to grow new anters in iate Apri and/or eary May, athough some carry od anters through to Apri or May; (3) many nonpregnant barren-ground femaes begin gro "' ing anters as eary as Apri, but some carry ast year's anters unti May or June~ (4) many pregnant barren-ground femaes stan anter growth about one week after caving. and, whie many carry od anters to caving. some cast their anters as eary as March or Apri. Hooves. The hooves of caribou are we adapted to ife on spongy muskegs and frozen snow-covered ground. The hoof is as wide as or wider than it is ong; dew caws are arge and add greaty to the bearing surface of the foot. The hoof is curved inward and abrupt at the tip. n winter, the soft pads of the hoof deteriorate and the pad area becomes concave. The hoof becomes sharp edged and hair between the hooves eongates to cover the pa.d. Those change,~ resut in a spreading, ''nonskid'' support that is effective on snow and ice (Kesa 968). A poydactyous hoof was found on a Canadian barren-ground caribou (Mier and Broughton 97). Sku and Dentition. Banfied ( 96 :26-27) diagnosed the sku characteristiqs (figure 47.2) of Rt.tngife r as foows: ''Cranium moderatey expanded, extr'!me postorbita position of the pedices which encroach on the parieta bone, nasas prominc:mt and exp'ndcd proximay. preorbita pit in the achryma bone moderate in size. achryma vacuity generay moderate in size. Premaxiae prevented from reaching nasas by a sma obe on the maxiae. Caribou are heterodonts (Loomis 925: Frick 937: the permanent dentition is subhypsodont. seenodont. and deerike in genera. but seemingy more adapted for grazing than for browsing {Kesa 968: Skoog 968). This adaptation of the teeth ha!.. however. been questioned by Mier (974cJ in th( ight of the known varied food habits of caribou. Tht denta formua is 0/3,. 3/3. 3/3 = 34. Neonata caribou have a set of functiona decidu ous mik teeth that are smaer versions of their perma ~ j \ FGURE Sku ofthe caribou (Rangifer rarandus). Fi om top to bottom: atera view of cranium, atera view of mandibe, dorsa view of cranium, ventra view of cranium, dorsa view of mandibe. _j nent countej the first mvi and the sec~ moars and 974c). W vear of if~e ~ima is 5 ciabe on their usefu. Denta. iations. s 954a. 9 anomaies by trauma Skin Gan digita. an materia ( 968~ An 978). Th eye socke bone. The' hair. A tar inside of. fringed by q yeowish' t.' g{uhds oc 'each fron!: digits. Th<\ ta portiorr Thes 'j roe in S<j: i..' rated me. of each o.i seem gaf! tory sign" different : each oth, as indiviq, track eac~ Longevi4 ates. Sog caribou~! 968), ( PHYSid Nutritio caribou '7 the adap environ main wi carbohy. Whitehe cay nd 975b).r,.. ow co. ;~i~~tj: pasma. cycina : 0 ' the o...,. t --=-= ~ ~ ~ ~ r"'"''"-'""-""""" - :~ ~ '-:cc'"') ~ ['

6 J ~-,~,,, i) t: 'Jf ~~:-' _,,, ;o'! uchrym.~ ::' yener..{- ::- '.. ',.m reachm"",925: Frid., hypsodom. :_t ~eeminf.' ing (Kesa.;e teeth ha!>. : 7 -k in tht ;;aribou. Tht./ 'pna decidu )heir penna i ~ f:>~ ft fi ~- ~ :andus). From : ew of.mandirnium, don;a nent counterpans. 'fhe first p'!nna,nent tooth t0 erupt is the first moar at 3-~ month'. foiowed by the incisors ar.d the second moar at 0-5 months. then the premoars and third mo;a. at month Mier 97;., 974c). Wear on the teeth is sight unti after the third year of ife. Funher attrition is n<t marked unti the anima is 5-7 years od: b?" 0 v.,.:.(<; the wear is appreciabe on most teeth and th~ rr.:,iars :ppear to be osing their usefuness. Dentai arwma!ies inc!ude morphoogica variation~. supernumeraries. a~ i agenesis fbanfied 954a, 96'?viier and Te".si~r 97). Many denta anomaies art: : ongenita. whie others may be c~used by trauma and subsequent rep~it Skin Gands. Caribou have preorhita. tarsa. interdigita, and cauda (tai) gands that a secrete odorous materia (Quay 955; Lewin and Stefox 967; Kesa 968; Anderson et a. 975; Muer-Schwarze et a. 978). The preorbita gands are ocate<~ media to the eye sockets, over the achryma pit in the achryma bone. They are amost hidden by a den.$e covering of hair. A ~arsa gand i~ ocated on each hind eg, r n the inside of the hock at the tarsa joint: the gands are fringed by tufts of stiff, whitish hair. usuay st' ~d yeowi 'h by the secretion. Deep-socketed nterdtg;.;! gands occur on each hind foot, and a sm~uer one ta each front foot, media and postedor to the primary digits. The cauda gand is ocated utr.;erneath the dista ponion of the tai. These scent gands most ikey pay an i!nponant roe in socia organization. The fact that c ;f.'\jou can track each other by scent makes it possibe for separated members to re~ate the group if they ose sight of each other. Future work wi undoubtedy show ~hat scent gands function in many ways to provide ofactory signas to caribou und.er various conditions and in different situations. f irrdividua caribou can track each other. why shoud they not recognize each other as individuas. rather th.~n by sex and age cass ony? Longevity. Caribou are moderatey ong-ived unguates. Some extremes of ongevity for North American caribou are 8-20 years (McEwan 963). 5+ (Skoog 968), 7 (Bergerud 97c), and 7+ (Mier 97""'). PHYSOLOGY Nutrition. Physioogica studies of nutitton of caribou are beginning to unrave the mechanisms of the adaptation of caribou to their arctic and subarctic environments. Lichens (Ascomycetes), usua!y the main winter forage, are ow in protein and high in carbohydrates (Scatter 965, 972; McEwan and Whitehead 970). The atter is fermented by specificay adapted rumen microorganisms (Dehority 975a, 975b) and during its fermentation heat is produc~d. A ow concentration of rumen ammonia aso resuts, which assists the recycing of nitrogen by increasing the rate of transfer of urea to the rumen from the pasni~ (Waes et a. 975). The increase of urea recycing in winter (Waes et a. 975) compensates for the ow protein co ttent of ichens and aso has a CARBOU ( Rangifer tarandus) 927 marker effec;t on the water fux (White 975). The tota amount of body \\'ater increases principay because w:.ner repaces decreasing body fat (Cameron and Luick 972: Cameron e~ u. 975J The increased voume of body water coud function as a themm buffer. especiay to the temperature changes caused by food and snow intake ccamevn et a! The decrease in nitrogen intake and the incre::se in urea recycing are aso accompanied by increases in gucose resynthesis CWhite!975>: the abiity to have a high rate of gucose resynthesis is particuary important in pregnant or actating femaes (Luick and White 975). The reduced forage intake during winter ( McEwan 968}. probaby in fespchse to ower energy requirements (White 975). in itsef reduces energy expenditure. Thing ( ) cacuated that the energy t.:osts of foraging in snow increase markedy in ate winter. By eating ess. the caribou saves the energy reqyired both to find food and to warm it once eatenan additiona cost estimated tc be percent of the fasting metaboic rate (White and Yousef 974; White 975; Young and McEwan 975). Caribou conserve energy in winter through the insuating yuaities of their peage and by heating the bood returning from the dista joints of their egs with outgoing bood ii\ the proxima anerie5 (Jacobi 93: Schoander et a. 950a, 950b. 950c: rving and Krog 955; Moote 955: Lentz and Han \ 960~ Han et u. 96; ~ritsand 974). The feet of caribou remain texibe in subzero temperatures because the fatty tissues formed there remain soft. Marrow fats further up the eg are soid even at room temperature. n summer, caribou ose heat by panting, from patches of bare skin on their bodies. and from the sp<rsey haired extremities (McEwan et i. 965~ rving ~:966: Krog and Wika.975; Yousef and Luick 97,. Stonehouse ( 968) suggested that the growing anters in vevet served a primary function in the dissipation of body heat. However. Krog and Wika (975) ~eheved that heat oss from the anters was unavoidabe because. :Pf the bood fow necessary for rapid anter growth. Growth. Caribou cai es grow rapidy in thf: first five months after binh. Their body weights increase by an average of 800 percent (Dauphine 976), from about 6 kg to 48 kg. Variations in body weight may refect the condition of the cow in ate gestation: nutritionay stressed cows gave birth to sma caves that had ow surviva rates (Skoog 968). Caribou normay grow between Apri-June and October-December of each year (Dauphine 976). The rate of growth depends on their nutiitionn pane in reation to other demands such as hair and anter growth reproduction, weather. and movements. incuding responses to insect harassment (Skoog 968). The age at which skeeta growth stops depends on which partic!ar bone is measured (McEwan and Wood 966; Daupiine 976). Mandibes continue to e.gthjn unti 5h years of ag~ in cows and 6 years in bus (Mier and McCure 973), but body weights reach maximums at 4h and 6 years for cows and buhs f

7 928 ARTODACTYLA respectivey (Dauphine 976). There are marked seasona fuctuations in body weight because of changes in fat deposit~ (Dauphine 976). Femae caribou gain \Veight annuay during ate summer through eary winter. Then their weights stabiize or decine in winter and are owest during eary summer due to pregnancy and actation. Adut maes gain the most weight from summer to the beginning of the rut. Marure prime maes typicay ose up to 25 percent of their weight during the rut: in winter their weight stabiizes or decines tdauphine 976J. Young animas reach maximum weights between eary summer and eary winter. Their weights decine during winter, as they usuay ack sufficient fat reserves to maintnii body weights unti spring. REPRODUCTON Caribou rey on high-quaity forage on their summ.er range for their reproduction, growth, and winter surviva. Femae caribou with ow fat reserves in the autumn do not breed but buid up their fat reserves and breed the foowing autumn. Body size and fat reserves of femaes are aso reated to the age of first conception: athough most cows conceive at 3V2 years of age, a few wi conceive at h years of age in good condition (Dauphine 976). This varies among popuations: more Aaskan than Canadian caribou conceive at eary ages (Skoog 968). Caribou are reativey sow to mature and do not bear twins, wit rare exceptions (McEwan i97; Shoesmith 976). The adut femaes, however. are very fertie, with pregnancy rates of about 80 percent or more. The chief cause of reproductive faiure is a faiure to conceive. as in utero mortaity is rare (Dauphine 976). nsufficient buid-up of fat reserves in summer and subsequent faiure to conceive may expain the fuctuations in pregnancy rates of some Peary caribou (Thomas and Broughton 978). A though femaes can be above the threshod of physica condition necessary to conceive. subsequent deterioration of the cow's condition can contribute to caf mortaity at birth or shorty thereafter. Death of the caf soon after birth reieves the cow of the metaboic cost of actation. This reief aows it to buid up fat reserves sufficienty to breed again the foowing autumn. but sometimes not sufficienty to produce and rear a caf successfuy. As a resut. Dauphine ( 976) suggests a direct reationship between high caf mortaity and high pregnancy rates. Breeding and Gestation. One of the most striking features of the reproduction of caribc'u is the synchnmizatkm of the rut and thus of caving (Dauphine ~nd McCure 974). Caribou have two or three fu estrous cyces, and one or more of those resut in ovuation without overt estrous behavior. Such "sier.t heats" coud generate and synchronize the endocrine system and thus mating (McEwan and Whitehead 972; Bergerud 975). Athough there is v.u-iation in the dates of rutting between different caribou popuations~ the rut is usuay in October and November, and oca synchrony of breeding is apparent. The average gestation period is days according to Skoog (!968). or day!> uccorju~ Bergerud ( 978). and the synchronized breed tn.:!>uh!- in a strongy peaked distribution of birth~ a) be~ween 80 and 90 percent of caribou cah c' born in a 0-day period in ate May or ear.~ Ju: Dauphine and McCure 974: Bergemd 975 L Parturition. Parturition has been observed by :-.e\,:: workers (Kesa 957: de Vas 960: Lent tjof. Bergerud 974d). The newborn caf"usuay stand",,; waks within the first hour of ife and nurse~ -.iw thereafter. The mother vigorousy icks and ceun'> u. neonate. She nibbes and pus the afterbirth fron around the caf and often eats it (Pruitt 960a; Mik and Parker 968). Mier and Parker ( 968) suggested that materna caribou frequenty consume the afterbirth. There ar two possibe reasons: ( ) remova of the tissue reduce" odors associated with the birth site and, thus. the chances of predation; and (2) the materna femaes derive nutritiona benefit from eating the afterbirth. The atter supposition seems most tenabe for caribou. Neonates foow their mothers and the birthpace i!. usuay abandoned within hours. Aso, the attending cow woud be sighted by a predator ong before tht predator was cose enough to detect the odors of tht. birth site, especiaiy on tundra caving grounds. ECOLOGY Habitat. The geographica distribution of caribou encompasses two of the argest biomes in North Am~ica: tundra and taiga (and their southern extensions in mountainous areas). Variations in cimate. geoogy, and topography from Aaska to Newfoundand resut in differences in the pant communities of these biomes. but there are aso simiarities in habitat types used by caribou. The typica range of eastern woodand caribou is cimax stands of northern borea forest with mixedage stands of back and white spruces (Picea mariana. P. gauca). basam fir (Abies basamea). and white birch (Betua papyrifera) with tree and ground ichens (Cringan 957).-The habitat of many western woodand caribou often incudes mourjtain summits above the timber ine or eevated tabe ands with apine meadows and open subapine forest (Edwards 958). n the taiga toward the treeine the trees become more widey spaced and the ground cover of ichens. ericaceous shrubs, wi!ows (Saix spp.), and dwarf birch (B. g/anduosa) increases. North of the treeine the back spruce-moss muskeg and sedge (Carex spp.) bogs of P-QQry drained area~ are repaced by wiow and aider (Ainus spp.) thickets aong watercourses, and by sedge in grassand communities on wet areas, Dwarf shrub-heath communities occupy arge;.areas of the tundra except on drier sites, where fructose ichens and mosses dominate with dwarf wiow, birch, and rhododendron (Rhododendron apponicum) (Rowe 959; Mier 976), n the High Arctic, Peary caribou show a preference for poar 'desert'' and simiar dry to mesic range cypes with sparse vegetation of wiow, sedges, grasses, and forbes (Parker and Ross 976; Russe et a. 978). area. forage a ing and often 959: H 975: M The partiay creased d. atitudes

8 t, ::---- CARBOU (Rangifer tarandus) 929 :cording t' eec.jing r~..,' rth:-..._,,. ', -:aj VC!\ ~tn: ~ary Jun~: :75. hy severa nt 966a:.': :-tand::. and : -.::~ shon: ceans the jjirth from jda; Mier., j r materna A There are i.te reduces thus. the!maes de, birth. The jr caribou. ~'thpace is [/ attending J : before the ors.of the ~~ nds. aribou en : America: :nsions in geoogy. d re<>ut in e biomes, s used by d caribou th mixedmariana. md white d ichens 'rn woodtits above ith apine is 958). s become f ichens, nd dwarf e treeine m:x spp.) y wiow!rcourses. vet areas. e areas of se ichens irch, and ) (Rowe y caribou iar dry to f wiow, >SS i976; The cimate of caribou ranges is ch:uacterized by ong. cod winters: short. coo summers: and ow precipitation. Much of the precipitation fas as snow. which cnn cover the ground for seven to nine months. Snow cover data are of particuar importance to understanding caribou bioogy but show great variation according to ocaity. The caribou contend with snow depths usuay exceeding 50 em except in the High Arctic and on the open tundra. Autumn and winter thaws and rain storms can occur in amost any northern area. These ~ometimes cause serious restrictions on forage avaiabiity (Mier et a. a). Spring meting and refreezing of snow eading to crust formation often restrict caribou feeding and movements {Pruitt 959; Henshaw 968b, Bergerud 974b; Stardom 975: Mier 976; Mier and Gunn 978). The short. growing. season on caribou ranges is partiay offset by the increased soar radiation and increased day ength characteristic of high atitudes and atitudes (Kein 964). n addition, pants growing on the ranges have higher nitrogen«phosphorus, and carbohydrate eves than pants at ower atitudes (Chapin et a. 975). Some arczk pants are adapted to the short growing season by having wintergreen and evergreen. eaves (Be and Biss ), which are preferred winter forage (Kesa 968). Caribou sometimes expose, trampe, and eat feeding push-ups constructed on ice in winter by muskrats ( Ondatra <.ibethicus) (Skoog 968; Kesa i970). Caribou wi use minera icks (Caef and Lortie 975). Summer and Winter Ranges. Most caribou popuations have distinct summer and winter ranges; the atter are chara-;tf.rized by tree and/or shrub cover and ichens. Caribou are not soey dependent upon ichens in winter and may take other forage if ichens are unavaiabe (Murie 935; Skoog 968; Bergerud 972). Wimer range!~ usuay regarded as the imiting factor for caribou popuations; therefore, any factors modifying or reducing rhe avai~abiity of winter range are of particuar interest to widife managers. A Peary caribou remain on tundra ranges throughout the year. Some barren-ground caribou herds remain a year on the tundra, whie other herds move t~:> forested ranges in winter. Woodand caribou, even the migratory herds of the Ungava P~ninsua. usuay move to forested winter ranges. However, even the forest-dweing herds wi move in winter to tundra or tundraike areas if excessivey deep snow cover restricts forest forage suppies. Movements between the tundra and forested ranges are earned traditions of particuar herds that ha ~; resuted in fayorab~ ~mg=t~rm ~B~rvi'gJ. Th~ yeary differences in movements refect the annua variations in snow c;over and icing conditions, which infuence avaiabiity of forage suppies. The effect of forest fires on winter ranges of caribou has ong been a controversia subject. There are two L'cories concerning the importance of forest firei.i in imiting or reducing numbers and changing distributions of caribou. One is that forest fires are a paramount consideration because of the extensive detrimentpj impact on forage suppy and the subsequent restorations of such stocks (Leopod and Daring 953: Edwards 954: Cringan 957: Banfied and Tener 958: Scatter a: Kesa 968J. Other theories suggest that ony certain fires under certain conditions are important to caribou. Finay. some researchers fee that fires are not ony beneficia to caribou hut aso are necessary to promote hc heterogeneity of vegetation needed to perpetuate mxed forage suppies Skoog 968: Bergerud 97 a. 97! 974a; Rowe and Scatter 973: Bunne er a!. 975: Johnson and Rowe 975: Mier 976L Probaby. the moderate views better refect the reaities of range caribou reations. athough production and avaiabiity of forage are the utimate factors in governing numbers and distributions of caribou. Therefore, managers must remain aware that wi!dfires can be important in the ecoogy of caribou. f rates of burning approach the time required for rotation of successiona ranges of vegetation to provide sufficient forage suppies for existing and future numbers of caribou, protective actions shoud be taken to minimize destruction of ranges. Movemefts and Migrations. f one aspect characterizes the ecoogy of caribou. it is surviva through adaptive movements and migrations. Bergerud ( 974d) beieved that interaction with woves ed to gregarious behavior. Movements and migrations foowed as a resut, so that caribou coud maintain themseves in reation to their varying forage suppies. The caribou's movements and migrations are further governed to varying degrees in time and space by weather, especiay snow cover and icing conditions, boodsucking and biting insects, and various physioogica and psychoogica drives. Snow cover most often is the dominant infuence on movements by caribou during most of the year. Evauations of the snow (niva}) environment of caribou have been made by many workers (Banfied 949: Pruitt 959~ Edwards and Ritcey 960: Henshaw 968b; Bergerud 97 a; Stardom 975: LaFerriere and Lent ; Thing ). The annua cyce of movements of migratory barren-ground caribou begins in spring. As the amount of dayight increases and snow begins to recede, wintering bands of caribou begin to coaesce, and these arge aggregations move northward. n the eary stages of the spring migration, the caribou move sporadicay unti the apparent urge to return to the caving ground grips the parturient cows and they move steadiy north. The number of juvenies, and especiay yearings, ac ~ompanying the femaes to the caving ground varies from year to year depending on the difficuties of migr;!!i<:>n. Jn SQf!~ Yf!~S! df!ep snow and sush prevent most young animas from migrating with the- cows. They drop behind and move northward with the bus at a more eisurey pace as traveing conditions improve. Once the traditiona caving ground is reached, the parturient cows disperse. They may cave in reative isoation or cose to groups, depending on the timing of their arriva on the caving grounds and the peak of caving. The materna cows and their newborn caves form nursery bands after the peak of caving (Pruitt 960a).! [

9 ARTODACTYLA The nursery bands then merge into arge postcaving aggregations that move off the caving ground to summer ranges and mix with the bus and the rest of the herd. During the summer months they move extensivey, often 500 km or more. Some herds return sot.:th into the borea forest. whie others move northward to tundra and coasta areas. n autumn, there are prerut movements and the prime bus estabish their dominance hierarchies. With the beginning of the rut. bus join cow-juvenie groups and remain in their company unti the cows become receptive. Breeding often takes pace whie the aggregations are on the move, sometimes even during the autumn migration to the wintering gn..:... :ds (Henshaw 970). After the rut, the caribou move to their winter ranges. Adut bus often separate from the cow-juvenie groups. and some groups of bujs move farther into the borea forest. Se'!--;!:!>:a movements of the more sedentary forest-dweing woodand caribou and Peary caribou are sma scaed by comparison with those of migratory barren-ground and woodand caribou. Loca aterations in movement patterns of some woodand and Peary caribou refect seasona variations in distribution and avaiabiities of forages,, and specific physica differences among the habitats. On western ranges, where the eevation can vary by over 2,000 m, the habitat infuences seasona movements of woodand caribou (Edwards 954, 958; Edwards and Ritcey 959, 960). The 25 to 30 remaining woodand caribou in the Sekirk Mountains of northern daho. northeastern Washington, and southern British Coumbia have stopped making distinct seasona shifts in eevation (Freddy 979). Woodand caribou on mountainous western ranges are found on high meadows and in adjacent, open subapine forests above 2,000 m during summer and fa. The first deep snow of winter forces them down to mature forests in vaijeys. where they prefer poory drained sites interspersed with open bogs, meadow~. and ponds. As winter progresses and the snow settes. hardens. and becomes crusted. the caribou return to their summer haunts on the high, wind~swept ridges and remain there unti spring. Softening snow drives them into owand forest again, where they remain unti they can return to the snow-free upands in May and June. Most, if not a, Peary caribou make migrationike trips at about caving time. They may rrave between isands, or around or across one isand. Movements are not necessariy we in advance (days or weeks) of c~hving, as they are with woodand and ban en-ground caribou (Mier et a. a, b; Mier and Gunn 978). Peary caribou seem to prefer certain area!i of different isands for caving, but to date no spe.:cific caving grounds are known. The Peary caribou constanty move and feed on their summer ranges unti eary fa. There appears to be a prerut movement, or ''fa shuffe," in ate August to September. Nothing is known concerning the rut in Peary caribm~. Cast mae anters found aong coasta areas suggest that there is some preference for the coast during the rut or shorty thereafter. This woud facii~ tate mae-femae contact. Herds that use more than one isand annuai: probaby return to winter range shorty after freeze-ur in ate autumn foowing the rut. The rut most ike!: occurs on the isand where the caribou summered. Peary caribou that remain on the same isands yearong shift from summer range to winter ranges on different parts of the isands (Freeman i 95: Mier et ui. a). Even the sedentary individuas must sometimes move to adjacent isands to survive when deep snow reduces forage avaiabiity on arge areas of their home ranges. Bioogists have described ca.ribou movements and migrations as nomadic (Skoog 968: Bergerud 974d). That description shoud not connote herds of caribou wandering. aimessy about tundra and taiga without purpose. Their wanderings are structured and carried out in an ordery sequence (Heape 93). Caribou herds, or segments of herds, prefer specific sections of ranges: not ony caving grounds but aso wintering areas; spring, summer, and fa staging areas; rutting areas; and migrationai paths. To maintain such traditiona range use requires a refined state of Ortstreue, or fideity of the offspring to the and of the parents. The apparent wanderings of individuas and groups of caribou are strongy orientated and directed. Wise management of caribou herds demands that the movements and migrations of caribou be studied in detai to determine if individua caribou show strong seasona affinities for specific sections of their ranges, f different caribou from one segment of a herd return to the same areas every year, continued heavy annua harvest of them woud not be detrimenta (assuming that the popuation coud sustain it), because different individuas woud be fiing the voids each year. f. however, individua caribou did show strong affinities for specific sections of the popuation's ranges, continued hea\"y annua harvest of them woud soon markedy reduce or destroy that segment of the popuation. Herd Composition. Caribou are highy gregarious. The zenith of their gregarious behavior is reached in t':e postcaving herds composed of tens of thousands of barren-ground caribou. Those herds are truy ''iving tides fowing over the arctic prairies." No one can ever forget the sights and sounds of such a mass of moving animas. Thousands of grunting cows are answered by their be<j.ting caves. There is a continuous cicking of hooves, and a myriad of coughs, sneezes, and beches coming from the mvving mass. Such aggregations sometimes bunr h so tighty as they mi about that from the air they.appear ike a swarm of bees. Large herds of migrating caribou are temporary gatherings of many socia units (groups or bands). Some socia order may be maintained within aggregations by an interacting hierarchy of do~inant animas from the bands that form the aggre;;.:.=~:s. Socia Structure. Caribou occur in different groupings, but consider the band the primary distinguishabe socia unit. characterize bands of barren-ground caribou by their representation, by sex and age compo-

10 CARBOU (Rangifer tarandus) 93 "'ttnd ~~:~~;~. i :mer tr". -.!-th fnp..,;l\l '\Unit!.~~ _.and..,~ C<r;"L e~ on~;!:~...-..,! :-,... fier ~.:,s mu!>t, 0 m ;ive when ke; '~area... n th"' ovemem.., 4 : ~erud 974tiJ rds of carihou taiga withmu.;=d and carn,: 93). Caribou ~ific section!'> oj ~o winterin! ' - fr areas: rutting use requires : offspring w jhe,twanderings o.wrongy orien- ~ t de~ands thttt u be studied in J show stronu. rf th~r range:-: a herd return heavy annua tta (assuming ause different tach year. f. ong affinities ranges. con- id soon mark e popuation.! ~ -gr~garious. S reached in.f thousands of truy "iving ) one can ever ass of moving : answered by us cicking of ;. and beches aggregations bmjt that from "e temporary JS or bands). ithin aggregainant animas JS. ~rerent group Y distinguish JarreR-ground d age compo-. ~ J i {. J i... smon: cow. bu. subadut. juvenie. cow-juvenie. cow-juvenie-bu. and bu-cow-juvenie bands (Mier 974c). Lent (966a and Bergerud (974d) considered caribou groups as open socia units with no ongterm stabiity. suggest, however, that at east the cores of barren-ground caribou bands represent cosed or semicosed socia units. There is evidence of group cohesion and eadership (Mier et a. 972) and of ong-term socia bonds in barren-ground caribou on the mainand of northern Canada (Parker 972b; Mier et a. 975a). f ong-term socia bonds do exist among members of a socia unit, one must consider how group unity is perpetuated from one year to the next. The basic socia unit of barren-ground caribou is the winter band. of which there are four types: bu, cow-juvenie. juvenie. and subadut. The juvenie and subadut bands are usuay mixed or in cose associatio~ with the cow-juvenie bands. Juvf"nie and subadut bands may be the resut of the oss of materna cows. Their ocations with respect to aduts may be governed by the antagonism of the remaining mature cows or dominant subaduts. The core of the winter cow-juvenie bands is formed by cari~ou reated through a matriarcha boodine, suppemented occasionay t;y neighboring cows and their young or juvenies, The vinter bu band recruits most three- and four-year-od maes from subadut groups in the fa, during the prerut period. Most band members have simiar body and anter size. Athough anters may vary, they are not usuay as arge as the anters of prime bus. Th.e bu band maintains a nu~..eus of breeding bus from year to year with common earned behaviora habits. Under norma conditions, this assures a suppy of breeders in the traditiona rutting areas. n spring, the bonds between members of the cow-juvenie bands weaken as the caribou bands come together and the caribou move north to the treeine. On the tundra, the panurient cows intensify their antagonism, and many of tie juvenies and subaduts drop behina. This response is reinforced by deep, wet snow aong the migration route, which sows the young animas. After their caves are born, the cows move about the caving ground and join other cows, caves, and those juvenies and subaduts that have arrived. Sma groups are attracted to arger groups unti, finay, they form postcaving herds of severa thousand caribou. A postcaving herd has a distinct structure: at the core are the materna cows with their caves, equay spaced throughout and occupying an eiptica or round area. Aong the periphery are groups of yearings and juvenies in constant motion. Some caribou rejpin feow members of previous winter bands during formation of postcaving aggregations. Large numbers of juvenies and subaduts eft behind may rejoin the cow component of their winter bands when the postcaving groups move into the area occupied by those young animas and mature bus that have moved northward after the cows. The cows and caves merge with the bus, remaining juvenies, and subaduts, and move off on their midsummer migration. The animas con- tinuousy shuffe and reshuffe. By the onset of rut the winter bands have reformed. They remain generay in arger groupings unti arriva on the wintering ground. The bu bands separate from the cow-juvenie bands and move off to their particuar wintering areas. beieve that socia!ization during the postcaving aggregations eads to the regrouping of previous winter bands. This ink is necessary for perpetuating beneficia behaviora traits (Mier 974c). which is of particuar concern to managers of caribou popuations. Barren-ground caribou usuay form arger groups and aggregations ~han woodand or Peary caribou form, athough some herds of migratory woodand caribou-for exampe, the George River ierd, in Quebec-Labrador-might be as arge during popuation highs. The sizes of groups refect the overa number of caribou in the herd, the size of their ranges. the dispersion of avaiabe forage, and possiby subspecific differences in behavior. f the groups are more or ess "cosed" socia units, their sizes woud a:;o depend on the surviva of group members. Group sizes coud vary consideraby (usuay 5-30). but coud sti incude a core of reated animas. The smaest groups of caribou usuay occur in midwinter, except for cow-caf pairs during caving. The sma winter groupings probaby refect the distribution and avaiabiity of forage suppies and possiby a high degree of socia intoerance for nongroup members. Forest-dweing woodand caribou form reativey sma socia units, probaby because of their secusive nature and the mosaic pattern of their habitat. During ate winter, those sma units merge into sma aggregations, probaby because foraging sites and resting areas are restricted. The average sizes of Peary caribou groups are sma at caving time and then. as with the other subspecies, their groups increase markedy in size during postcaving movements. Such aggregations can exceed 00 caribou but average much ess. Winter groups of Peary :;:aribcu are sm!t-amost aways fewer than i 0 individuas, and usuay averaging ony 3 to 4 animais (Mier et a. a )~because forage is reativey imited on most of their wintering areas. nterspecific Reations. Most interspecific interactions between caribou and the other animas on their ranges are either predator-prey or competitive. Animas that do not interact directy with caribou benefit by scavenging upon them. The wof (Canis upus) is the principa predator of caribou, but on some ranges, especiay where woves have been reduced, the ynx tfeis ynx) and the grizzy bear ( Ursus arctos) are more important. The back bear ( Ursus americanus), the woverine ( Gu/o gu/o), the coyote (Canis atrans). the n;d fox ( Vupes vupes), the bobcat (Feis rufus), the goden eage (Aquia chrysaetos), and the raven (Corvus cora.x) sometimes prey on caribou, especiay nev.born caves. Most potentia competitors inc!ude muskoxen, emmings, (Dicrostonyx spp. and Lemmus spp.), arctic hares (Lepus arcticus), and snowshoe hares (L. americanus). Minor or theoretica ~ompetitors coud '

11 -! 932 ARTODACTYLA incude other un2:uates. other rodents. and grazing waterfow (Kesa 968: Skoog 968}. Scaven!.!ers other than the previousy named predator~ \Voud incude the poar bear< Ursus maritimus}. the arctic fox (A/opex agopus), gus (Larus spp.), and jaegers (STerocorarius spp.). There are a_ host of other potentia scavengers. such as musteds and carrion-eating birds (Kesa 968: Skoog 968). Fv Season. Perhaps no other aspect of the caribou's reiationship to its environment is so vivid as its stressfu encounters with boodsucking and biting insect> during summer. A year when such insects are numerous is- indeed a time of madness for the caribou. There are no areas free of harassing insects on the inand tundra. Caribou must constanty dash about widy to escape the ever-present hordes of fies du~ng the diurna peaks of insect activity. Fortunatey, m ~ but the worst fy years, there is some temporary rehef for the caribou during the cooer eary and ate hours of the day. Caribou occupying coasta summering areas often seek reief from fies by moving out onto the mud fats and simpy standing motioness with their heads down, muzzes neary to the ground. Windswept ridges, gaciers. and ingering snow drifts on tundra ranges of some barren-ground caribou and most western woodand caribou and and areas above the timber ine are free of fies. On those sites, caribou usuay remain hudded cosey together throughout much of the day. dispersing to feed in the cooer ate and eary hours. n Newfoundand, Bergerud (974d) observed that woodand caribo? di~persed into the forest. because fies were ess active _m the shade. This option appears to be the ony one avaiabe to most of the woodand caribou of eastern North America, uness they are on coasta areas or the shores of arge akes.. Peary caribou on the more southern Jsands of the Canadian arctic archipeago experience fy seasons of short ~hmuiqn that woud s~!dom, if ever. have mu~h mpact on their physica condition. Fies in certain vears coud stress them, but this woud be of minima impact compared to mainand situations.. Ony the Peary caribou of the more northery Htgh Arctic isands are continuousy free of harrassment by insects. These animas might not survive the rigors of the High Arctic if they had to withstand the additiona stress of harassment by hordes of insects. FOOD HABTS Caribou ike most North American cervids, feed on a broad ra~ge of pants, incuding ichens, fungi, sedges, grasses, forbs, and twigs and eaves of woody pants. Their preference for ichens is unique ~mong N~rth American unguates, and is the key. to car,~ou surviva in many areas. Descriptions of fee~mg habits ar.e base_d on direct observations of free-rangmg and capuve ammas, and aso upon examination of feca _an~ rumen sampes, fistua experiments, and e~anunau?ns of craters and other feeding areas. Feedmg habjts and preferences vary according to range type. Much information has been compied for woodand caribou in Newfoundand!Bergerud and Noand 970: Bergerud 97 a). in Ontario ( Cringan 957: Simkin 965. and in British Coumbia (Edwards and Ritcey 960: Edwards er a. 960): for barren-ground caribou in Canada (Scatter 967b; Kesa 968: Mier ) and in Aaska <Skoog 968: Kein 970a. 97Gb: White et a. 975: Kuropat 978): and for Peary caribo: ("Parker and Ross 976: Wikinson et a!. 976: Parket 978: Thomas et a! : Shank et a!. 978). Mos\ caribou ranges are fragie and sow to recover from misuse. Many of those ranges have short arowine seasons with ow productivity. Aso, ichens :re esp~day susceptibe to tramping. t is unikey. however. that caribou woud destroy their own forage suppies (Skoog 968): () their cu:sory feeding be havior prevents excessive use of individua pants; (2) they seect mosty the newer growth pans of pants, which are readiy repaced; (3) they have diverse feeding habits and use a wide range of pants; (4) t!ey use primitive pant (ichen) communities; and (5) snow often covers arge areas of ranges causing caribou to move to other areas. The wide range of habitats and snow conditions that determines caribou feeding habits imits the vaue of generaizations. Nevertheess, some comments are appicabe to many caribou. Foiose ichens dominate their diet in the fa and winter and, in deep snow areas, the terrestria ichens of eary winter are repaced by arborea ichens as snow depth increases. Many woodand caribou in the western states and in western Canada rey on arborea ichens for winter surviva (Edwards and Ritcey 960; Edwards eta!. 960). Stardom ( 975) reported that woodand cuj."ibou in Manitoba fed mainy on arborea ichens in open tamarack (Larix /aricina) bogs during eary winter unti snow cover appeared to hinder trave. They then moved to mature pine (Pinus divarkatq) ic:htm ridges and fed mainiv on the ground ichens (C!adonia spp.). During spring the wo~dand caribou fed aong shoreines _on Carex spp. and on ground ichens on southeast-fa~mg sopes of rocky ake shores. Barren-ground canbou sekct fungi in fa (Kesa 968; Mier 976). The winter diet in.cudes wood igs of shrubs such as Vaccinium spp. and some! )afix spp.), evergreen eaves and graminoids that :retained some green t - eaves (Kesa 968). As the snow mets, caribou seek exposed sites to feed on eaves and graminoids. which tend to dominate the diet tp the excusion of ichens as summer progresses. n particuar, at that time of year, cari~ou seect pant species according to phenoogy of greemng eaf buds and fower buds. The seectivity is cosey tied to the nutritive status and chemica defense posture of the part of the pant (Kuropat 978), The caribou feed whie on the move, and tend to concentrate on one or two species at a particuar time (Kuropat 978). They use sight and sme to seect preferred pant parts (Wright, in Kein and White 978). Litte is known about how seectivey caribou feed during fa and winter, but imited avaiabiity may reduce seec di fferem i~ 969a. 9t! probaby de sense of 5m through air 5 ~,~rfrrce tbe shaow snc increa<;e th~ front feet r caribou rna threshods wind. surf rain-that n the taig caribou sto e/cm 2 {Prui ~ontinued 6,500-9,0 ) and a. ), - vary from ( BEHAVC Behaviora tory. and o tion. They pays of d, and ~ecret~ messages 'i (affin typ 974). The af tion in des(!;.' socia sign{ which antf Dur!more t Lem c 96 (973. 9! Pruitt brow tine "bush-thr.., swinging i ~ege.tation. une s use (bez) tine Hens anters by, ing and, stricted for viva and anters o~ shaw, on9 maes thaf' a of the associatio- and their~ popuatio.. not poss. (968a) t caves a

12 e ; sun iva e j ~ribou in i3er!!eruj L--,, 7v=' i. ~ O!!':oU: Eu /nbou in ;t!r c., ir'or PearY )al 976: \nk et a. J ' >w to re- 'hve short. ichem. 'unikey. ' n fn... no:pt '.4"''. "'"""~- ~dmg be- ~ant;; (2) pr pants, frse feed / they use fj5) snow raribou to,fionditions thhe vaue ~ments are jdominate :ow areas, pac~d by iny wood- western t60). Stc..,? in Man! tamarack tntij S\ JW jmoved to rs and fed.j. Dunn!!!reines ' on - ast-facing d caribou 976). The s such as evergreen Jme green ed sites to, dominate nmer pror. caribou f greening is cosey tse posture he caribou rate on one pat '978). piant parts reduce sekctivity. Captive caribou and reindeer prefer differem ichen species (Des Meues and HeyJand 969a. i 909h; Hoeman and Luick ). Caribou probaby detect the ichens under snow by their keen sense of smel n deeper snow they may sme ichens through air vents caused by shrubs reaching the snow's surface (Bergerud 974b). Caribou wi move soft shaow snow with their noses, but as snow depths increase they dig craters of varying sizes with their front feet to expose vegetation. f the snow is crusted. caribou may break the crust with their front feet. The threshods of depth and crust hardness-caused by wind, surface meting and refreezing, or freezing rain-that preven~ cratering vary according to habitat. n the taiga, where snow tends to be deep and soft, t:rr..ritxm ~tgp,_ Cr-!t~nns when snow hardness exceeds 50 g/cm 2 (Pruitt 959). On the tundra, however, caribou continued to crater unti snow hardness reached 6,500-9,000 g/cm 2 in Aaska (Henshaw 968b; Thing ) and 0,000 g/cm 2 in the High Arctic (Thomas et a. ). Critica snow depths for cratering by caribou vary from 60 em (Pruit! 959) to 75 em (Thing ). BEHAVOR B~uwiora messages can be passed by visua, auditory. and ofactory modes either singy or in combinati.on. They are manifested as acts of aggression, dispays of dominance, vocaizations, skeeta sounds, and secretions of pheromones. Supposedy, a these messages have the meaning of either "come coser" (affin type) or "move away" (diffug type) (Lent 974). The anters of caribou deserve specia consideration in descriptions of their behavior, as they have high socia significance. Behaviora posturing or gestures in which antk -; have a roe have been described by Dugmore \':3), Pruitt (960a), Espmark (964b), bent (965a;!965b), Hgnsh!!W (968a). Bergerud (973. t974c) and Bubenik (975b). Pruitt ( 966) beieved that the we-deveoped brow tine (shove) protects the eyes during bouts of ''bush-thrashing" (stereotyped movements of the head, swinging anters back and forth through bush-type vegetation). Bubenik (975b) beieved that the brow tine is used as an offensive weapon and that the second (bez) tine protects the eyes and facia~ region. Henshaw (968a) beieved that retention of anters by pregnant femaes enhances their socia ranking and, therefore, their abiity to compete for restricted forage suppies, thus cm:cributing to their surviva and that of the fetus. Athough the occurrence of anters on femaes is acceptaby expained by Henshaw, one might ask () why are one- to three-year-od maes that often retain their anters throughout most or a of the winter permitted sociay to remain in cose association with wintering groups of pregnant femaes and their young? and (2) why do many fettaes in popuations of caribou, especiay in Newfoundand, not possess anters (Bergerud 97a)? Henshaw ( 958a) aso beieved that possession of anters by caves adowed thetn to exhibit a marked degree of CARBOU (Rangifer tarandus) 933 precocity and independence from materna care during winter. Athough such behavior by caves is ikey necessary. as pregnant materna cows offer itte or no care. ther sma spike anters are probaby of itte significance in encounters. Cows are often even antagonistic toward their previous year's offspring during gestmion. e~pcciay in \vinter if forage hecomes ~~stricted. Femae caribou w\thout anters often attack by kicking with their foreegs. Vocaization and Other Sounds. Caribou of both sexes and a ages produce many sounds. espt:ciay when in arge postcaving aggregations and during the rut (Mutie 935: Banfied 954a: de Vos 960: Pruitt 960a; Espmark 964a; Lent 965b. 966b, 975; Kesa 968: Bergerud 973, 974c: and Erickson i 97'). Cows give.::oughike grunts and caves have beating cries: rutting maes grunt, snort, surp. cough, sneeze, and pant. When caribou move, their hooves make a unique cicking sound that is caused by the sesamoid bones sipping over each Nher. This noise is particuary noticeabe when a arge number of caribou are on the move and, in cairn wea~her. at a distance over 00 rn, even when mixed wich assorted vocaizations. Locomotion. Caribou have severa gaits, incuding a wak, fast wak. trot, pace, and gaop. They wak whie feeding and moving between feeding areas. Athough waking during maintenance activities seems eisurey, it bs been timed at 7 km per hour over ong distances and on rough terrain (Pruitt 960b). Fast waks often occur during migratii)t and periods of initia aertness. Trotting often foows <!n aarm or stresses such as predators or insect harassme~t. f the aarm or stress situation continues and/or increa~es, caribou may pace or. more commony, gaop. Caribou are adept cimbers and wi ascend ciffs and traverse gacia snowfieds. They cimb sheer was of hard paskgd sncw by digging stgp!_: with th~ir front hooves as they make their way up and over the barriers. Surprisingy, they often make those steep cimbs when there are pathways avaiabe that offer itte or no re:;istance Caribou are strong swimmers and readiy cross swift rivers and swim akes during their traves, again even when not apparenty necessary. Their broad hooves and dew caws act as efficient paddes. They swim with their heads hed high out of the water at speeds ranging from 3 to krnh (Seton 927; Banfied 954a; Kesa 968). Reproductive Jehavior. PRERUT. n September and eary October, the bus strip the vevet from their anters, become irritabe, and engage in sparring bouts with other simiar-sized maes to estabish socia rank. Bus that appear dominant sedom actuay fight subordinates, but effectivey use their anters and posturing to reinforce their status (Bubenik 975a). Other common breeding dispays seen during this period of sorting out sociay are: bush gazing (stand motioness, with a fixed distant gaze) (Lent 965b), bush thrashing. hock rubbing or tramping (take up a [! --

13 rtribou in J. j PerncruJ ') ft'-.:' ~- un!. "4 '> E. ~ a : u- ; : Jribou in, ~r 9:'-. : CJ70c,, r: or PearY a. 976:, nk et a. ; Jw tore. r Vf! short : i. ichens ' unikey. n fora2e. ' - ~ ding be-,,!ants; (2) ) f pants. J rse fe,~d,. they use, (5) snow, j-aribou to western surviva 60). Starin Mantamarack nti snow moved to s and fed.. Dunng reines on :ast-facing d caribou 976). The s such as evergreen :>me green ed sites to >dominate nmer pror. caribou ' ~~greening S cosey tse posture he caribou rate on one ; pat r978). pan: parts may ' reduce seectivity. Captive caribou and reindeer prefer different ichen specie~ (Des Meues and Heyk_d 969a. 969/J: Hoeman and Luick ). Caribou probaby detect the ichens under snow by their keen sen~e of sme. n deeper snow they may sme ichens through air vents caused by shrubs reaching the snow's!-~urface fbergerud 974h. Caribou wi move soft shaow snow with their noses, but as snow depths increase they dig craters of varying sizes with their front feet to expose vegetation. f the snow is crusted. caribou may break the crust with their front feet. The threshods of depth and crust iardness-caused by wind. surface meting and refreezing, or freezing rain-that prevent cratering vary according to habitat. n the taiga. where snow tends to be deep and soft. caribou stop cratering when snow hardness exceeds 50 g/cm 2 (Pruitt 959). On the tundra, however, caribou continued to crater unti snow hardness reached 6,500-9,000 g/cm 2 in Aaska (Henshaw 968b; Thing ) and 0,000 g/cm 2 in the High Arctic (Thomas et a. ). Critica snow depths for cratering by caribou vary from 60 em (Pruitt 959) to 75 em (Thing ). BEHAVOR Behaviora messages can be passed by visua, auditory. and ofactory modes either smgy or in combination. They are manifested as acts of aggression. dispays of dominance, vocaizations. skeeta sounds. and secretions of pheromones. Supposedy. a these messages have the meaning of either ''come coser" (affin type) or "move away" (diffug type) (Lent 974). The anters of caribou deserve specia consideration in descriptions of their behavior, as they have high socia significance. Behaviora posturing or gestures in which anters have a roe have been described by Dugmo:e (93). Pruitt (960a). Espm!!c (964b). Lent (965a, 965b), Henshaw (968a), Bergerud (973, 974c) and Bubenik (975b). Pruitt (966) beieved that the we-deveoped brow tine (shove) protects the eyes during bouts of "bush-thrashing" (stereotyped movements of the head, swinging anters back and forth through bush-type vegetation). Bubenik (975b) beieved that the brow tine is used as an offensive weapon and that the second (bez) tine protects the eyes and facia region. Henshaw (968a) beieved that retention of anters by pregnant femaes enhances their socia ranking and, therefore, their abiity to compete for restricted forage suppies, thus contributing to their surviva and that of the fetus. Athough the occurrence of anters on femaes is acceptaby expained by Henshaw, one might ask ( ) why are one- to three-year-od maes that often retain their anters throughout most or a of the winter permitted sociay to remain in cose association with wintering groups of pregnant femaes and their young? and (2) why do many femaes in popuatk:.-.s of caribou, especiay in Newfoundand, not possess anters (Bergerud 97a)? Henshaw (968a) aso beieved that possession of anters by caves aowed them to exhibit a marked degree of CARBOU (Rangifer tarandus) 933 precocity and independence from materna care during winter. Athough such behavior by caves i!. ikey necessary. a~ pregnant materna cows offer itte or no care, thejr sma spike anters are probaby of itte significance in encounters. Cows are often even antagonistic toward their previous year's offspring during gestation. especiay in winter if forage hecomes restricted. Femae caribou without anters often att::::::k by kicking with their foreegs. Vocaization and Other Sounds. Caribou of both sexes and a ages produce many sounds. especiay when in arge postcaving aggregations and during the rut (Murie 935; Banfied 954a; de Vos 960: Pruitt 960a; Espmark 964a; Lent 965b. 966b, 975: Kesa 968; Bergerud 973, 974c: and Erickson 975). Cows give coughike grunts and caves have beating cries: rutting maes grunt. snort. surp. cough. sneeze. and pant. When caribou move, their hooves make a unique cicking sound that is caused by the sesamoid bones sipping over each other. This noise is particuary noticeabe when a arge number of caribou are on the move and, in cam weather. at a distance over i 00 m. even when mixed with assorted vocaizations. Locomotion. Caribou have severa gaits, incuding a wak, fast wak. trot, pace. and gaop. They wak whie feeding and moving between feeding areas. Athough waking during maintenance Ut.;L;,-ities seems eisurey, it has been timed at 7 km per :0ur over ong distances and on rough terrain (Pruitt 960b). Fast waks often occur during migratic..n and periods of initia aertness. Trotting often foows an aarm or stresses such as predators or insect harassment. f the aarm or stress situation continues and/or increases, caribou may pace or. more commony, gaop. Caribou are adept cimbers and wi ascend ciffs and traverse gacia ~nowfieds. They cimb sheer was of hard-packed snow by digging steps...;.,ith their front hooves as they make their way up and over the barriers. Surprisingy, they often make those steep cimbs when there are pathways avaiabe that offer itte or no resistance. Caribou are strong swimmers and readiy cross swift rivers and swim akes during their traves. again even when not apparenty necessary. Their broad hooves and dew caws act as efficient paddes. They swim with their heads hed high out of the water at speeds ranging from 3 to krnh (Seton 927; Banfied 954a: Kesa 968). Reproductive Behavior. PRERUT. n September and eary October, the bus strip the vevet from their anters, become irritabe, and engage in sparring bouts with other simiar-sized maes to estabish socia rnnk. Bus that appear dominant sedom actuay fight subordinates, but effectivey use their anters and posturing to reinforce their status (Bubenik 975a). Other common breeding dispays seen during this period of sorting out sociay are: bush gazing (stand motioness, with a fixed distant gaze) (Lent 965b), bush thrashing, hock rubbing or tramping (take up a : r

14 - 934 ARTODACTYLA hunched position and in most cases urinate on the hocks) (Espmark 964a: Lent 965b). mock battes. rearing. and faiing. Rt.:T. The intensities and kinds of behavior exhibited during the rut vary according to subspecies. habitat. and possiby the nutritiona state of the breeders. Open habitat appears to be preferred. if avaiabe. Barrenground bus do not gaiher harems (Pruitt 960a: Lent 965b: Skoog 968). Dugmore (93) suggested that mae woodand caribou gather hare ns~ however, Bergerud ( 973, 974c) stated that such groupings are not true harems because members of such groups are often transient. The preiminary phase of the rut invoves testing the estrous state of femaes; maes drive femaes, who fee before them. A modified form of threat (figures 47.3 and 47.4) is often exhibited by the mae as it drives a femae (Pruitt 960a; Lent 965b). Pursuits of femaes by maes are often interrupted by fights with other maes. n contrast to sparring during prerut, fights are often vigorous encounters (Lent 965b). Banfied ( 954a) reported mortaity from rutting fights and Bergerud ( 97 c) attributed high mortaity among prime bus to fighting during the rut. Maes aso stop pursuing the femaes to bush-thrash and bush-gaze. As the rut peaks, prime bus concentrate on tending estrous femaes and markedy reduce their foraging. The bu tends ony one cow at a time and foows it wherever it goes (L~;;nt 965b). Tending bus usuay show agonistic behavior ony when another anima approaches the tended cow. Tending bus tramp and bush-gaze. Ftav~g Top. outine and tona pattern of norma (eft) and threat (right) poses as presented by a cow to another cruibou ( Rangifer rarandus); bottom, outine and tona pat tern of threat (eft) and attack (right) poses as presented by a bu to another caribou. MoTHE its caf ing the icits bond. B visua. :taure Top, threat pose, exhibited by a fetnae caribou (Rangifer rarandu$): bottom, modification of threat pose by mae caribou. in courtship dispay. Copuation is rapid and has rarey been observed (Kesa 968). Bergerud (974c) suggested that the much greater weight of the mae requires that copuation be brief. E!.pmark ( 964a) beieved that for Swedish reindeer copuation occun ed ony once with each cow and mosty at dawn or dusk. CALVNG. On the caving grounds. the pregnant cows space themseves in smaer groups (maternity bands) (Pruitt 960a). n Aaska, caving groups were open and transient, but showed both intragroup and intergroup socia faciitation (Lent 966a). Femaes that drop their caves eary or ate tend to separate from other caribou, probaby because the groups remain on the move. Lent (966a), Kesa (968), Skoog (968), and Bergerud ( 975) a suggested that, during peak caving tim~. cows about to give binh do not activey seek isoation. Pruitt ( 960a) stated, however, that femaes giving binh during the peak of caving move off a short distance to cave and then rejoin their groups. The caribou caf moves off with the mother within hours after birth. The materna cow uses head bobbing (moves its head up and down on a vertica pane from the ground to about the height of the caf) {figure 47.5) and vocaizatior.s to strengthen the caf's foowing response (Pruitt 960a: Lent 966a). After the caves are mobie, "nursery bands" composed amost en-

15 CARBOU (Rangifer tarandus) 935.,.. rat (eft) fther, nu patfed by a copuiajhnt for ce with tire!;. of the caves and their matema cows are formed!pruitt 960u). MoTHER-YOUNG Bmm. The mother icks and feeds its caf within the first minutes of the caf's ife. initiating the mother-young bond. As the caf grows it soicits care from the mother and further deveops that bond. Bonding and individua recognition invove visua. auditory, and ofactory stimui. A strong mother-young bond is necessary for the surviva of offspring during the first six ftonths of ife. n the absence of a strong mothernyoung bond, most newborn caves woud die during th~ rapid and extensive postcaving movements of the arge herds. The motheryoung bond assures to a great extent that the materna caribou wi provide its young with ( ) passive immunity through coostrum and mik, (2) nutrition, (3) thermoreguation by direct contact and indirecty by icking and drying. (4) assistance in traversing difficut terrain, (5) defense against predators, (6) optimum environment for rapid deveopment and earning, and (7) behavior patterns that shied the infant from extremes of socia and nonsocia stimui (de Vos 960: Pruitt 960a, Lent 966a: Kesa 968: Skoog 968). After the death of a mother or its young. the sur ~iving member wi show a retention of the bond, sometimes for severa days (Banfied 954a; Lent 966a: Mier and Broughton 973). Cows with caves do not readiy accept other caves during the postpartum period (de Vos 960: Pruitt 960a: Lent 966a). Pruitt (960a) observed, however. that cows deprived of their caves wi accept strange caves, NURSNG. During the first two days after a caf is born, it nurses about every 8 minutes, usuay for periods of ess than minute. Caves over one week of age nurse ony about one-third as frequenty as younger caves (Lent 966a). The most common position for nursing is reverse parae. which aows the mother to ick and nibbe the caf's anogenita and back regions. Such care-giving behavior from the mother proongs the nursing period (Lent 966a). Caves aso nurse from the rear, sometimes whie the mother is waking. Litte is known of the actua weaning process in caribou, K~san ( 968) stated that sucking was greaty reduced by eary Juy. Biting insects woud greaty disrupt nursing after Juy. He concuded, therefore, that weaning must occur at about that time. Athough his concusion is based on considerabe observation, it is difficut to accept because of the high energy demands of the rapid growth of the caf during its first six months of ife. Skoog (968) suggested that caves are weaned between September and December, most ikey before November. The caf probaby associates. with its mother during most of the first year of ife, not ony because the mother is a source of high nutritiona energy but aso because the mother -young bond fosters psychoogica we-being. Fonnation of "peer groups" by short-yearings (ca.yes of that year) contributes to the deterioration of the mother-young bond and faciitation of the caf's assimiation into its mother's socia group. Agonistic Beha., ior. THREAT AND ATTACK. Pruitt C 960a) described and iustrated threat and attack poses by caribou figure -+7.6). He suggested :hat threat posturing (muzze extended. anters back) is used by femaes to ward off strange caves or aduts. by cafess femaes showing antagonism towaid maes during the caving period. :ind by ma!es to chaenge other mae5. Threat posturing aso is part of courtship dispays during the rut. and is used by dominant animas to defend or scare subordinates from feeding craters in winter. Attacks (head owered. anters presented) occur when the threat pose fais to intimidate or when the individuas are more high!y motivated. Attacks commony occur when two femaes compete over a singe caf. when juvenie animas are te\~~ng their strength or paying. and when two or more maes are sparring before or during the rut. DOMNANCE-SUBORDNATON, A system of rank order probaby exists invoving a individuas within a socia unit. Age, sex, size, and p'ossession of anters are a important criteria in determining rank order. Prime maes are usuay dominant over a other animas during the rut: oder, arger bus with bigger anters are J FGURE 47,5. Head-bobbing pose, exhibited by a materna cow to its newborn caf caribou (Rangifer rarandus). FGURE.,..6. Attack pose, exhibited by a mae caribou (Rangfer tarandus). ~ ~ ~ 3! ji ~ i " ~ i ~

16 936 ARTODACTYLA.' dominant over younger. smaer bus with sma anters. Subordinate animas are not aowed into the feeding craters of dominant animas in winter. Aso. domina~n caribou wi dispace subordinate animas from feedin!! craters. Materna cows are usuay dominant over other caribou in winter. Adut animas usuay dominate a juvenies (8-22 months od) and yearings (6-0 months od). As winter progresses. tht socia status of short-yearings changes as the motheryoung bond weakens. Lent ( 966a) argued that caribou recognize sex and age casses and that the dominance hierarchies are based on casses rather than on individuas. Parker (972b) and Mier et a. (975a). however. obtained evidence for ong-term associations between caribou, suggesting that caribou Q!.) indeed recognize individuas. Bergerud (973) suggested that the estabishment of a hierarchy depends on frequent reinforcement through interactions, but it may be necessary to have intensive interactions ony for reativey brief periods of the year. ALARM. Pruitt (960a) described and iustrated the unique aarm stance (one hind eg spread, head erect) and excitation jump (rearing.. pivoting, interdigita 2ands dischar2e). Both acts are highy noticeabe to ~tber members- of the group and, thus, warn animas that do not actuay see the danger. The aarm stance occurs when a caribou is aerted to some unidentified stimuus. The excitation jump usuay occurs after the caribou has investigated the foreign stimuus by sight or sme and has begun to fee. Low-intensity aarms incude head ow aerts (brief poses during feeding activities) and head high aerts (more intensive and proonged poses made by moving, bedded, or feeding caribou). Predator-Prev Behavior. The caribou has evoved with the wof: and undoubtedy much of the caribou's behavior has deveoped as a resut of that reauon::or.:p. The gregariousness of caribou, especiay after caving. probaby resuted from wof-caribgu encounters (Bergerud 97b, 974d; Mier 974c). The caribou's use of openings, frozen bogs, and akes for resting areas is aso a earned behavior that seemingy resuted from wof-caribou interactions. The caribou's apparent r.:~iugtance to enter riparian wiows and ot~er heavy brush, and its state of aertness when passmg through, suggest that it associates such cover with attack by wove(' and bears. Caribou have earned to distinguish between hunting and nonhunting wo :es. The- deveopm~nt of specific fight reieasers (Pruitt 965) and threshod distances aows the caribou to conserve energy by not taking unnecessary e~cape during wof-caribou e~cmmters. POPULATON DYNAMCS Reproduction, mortaity, and movements ( emigrations) reguate the number of caribou within the popuation. Mass emigmtions are not predictame and it is not known why they occur. Dispersa may be density reated. that is. a threshod density triggers socia intoerance and eads to disper!>a of!>orne.segment of the popuation. t is difficut. however. to perceive how a density concept works. because the cumping behavior of caribou on winter feeding sites and postcaving aggregations suggests thm caribou are toerant of high densities. Basicay. a popuation increases when recruitment exceeds monaity during a time period: it decreases when the opposite occurs. Nataity. The feniity and binh rates in caribou are usuay about the same from year to year (Skoog 968~ Parker 972a; Bergerud 94a: Dauphine 976). However, in some years disease or poor nutrition can cause high intrauterine monaity in some herds (Skoog 968: McGowen 966; Neiand et a. 968) and the birth rate drops reative to the ferti'ty rart~. The number of femaes reaching sexua maturity governs the potentia rates of nataity within a popuation. Free-ranging caribou caves are sedom bred. t is unikey that femaes bred as caves coud produce and rear young successfuy, because they are physioogicay and, probaby, psychoogicay unsuited for the task. Even the yearing femaes in many popuations are rot ready to be mothers. They usuay ack the necessary fat reserves and they are sti growing during the second winter of ife. Aso, young, primiparous mothers may ack the psychoogica adjustment for caving, and the subsequent mother-young bond either does not deveop or is weak (Skoog 968; Mier and Broughton 974). Most f~maes have to ive at east three years before they successfuy produce and rear their young. Athough the reproductive rate fuctuates ony sighty, annua yearing increments to the popuation often fuctuate markedy. Most of the information on mortaity in Rangijer has been obtained from studies of reindeer because of commercia interest and the reative ease of investigating semidomestic herds (Skoog 968). Forage. The utimate imiting factor on caribou popuations is theoreticay their maximum seasona forage suppies. Forage production and avaiabiity give a true approximation of the range's abiity to support caribou. Bergerud (974a) stated that a winter shortage of forage for free-ranging caribou has not changed binh or death rates in any popuations of caribou. However, data on those rates are difficut to obtain and the ack of evidence is not concusive that such changes have not occurred. Forage shortages occur in winter due to adverse snow cover and icing conditions. Forage produ.;: tion as a imiting factor woud be density dependent. t is unikey that other mortaity factors aow caribou numbers to reach such criticay high popuation eves on most ranges that forage becomes the controing factor. Weather. Weather, aone or combined with mortaity factors. coud hod a caribou popuation at a ow eve where density-dependent factors are not effective. Utimatey, caribou are governed more by precipitation in the form of " cod rains than' some de!!ree. s: if caribo~u ca~~ that!!reat\' re. : m.. ~emp~ratu suit in hi!!h ~ions of f~ragj sequent death f there is itte e ~ weather cond mainand. thei! on the High f beieve, as di~ attributed ro \ conditions. tt weather. not id MORT ALT' Caf Morta grounds is co t adverse weaq 968: Skoog[. Mier and B. wind are a c' often resutir respiratory p Broughton, snow traps c 968). Wear:. forage to pa; recty causes condition is ~ Canadian HJ Thomas andj Predati and often tr Broughton :! seek caves pacts on ca from popu(; from year tt ki n.:>stv ~ ' way. Atho! ame caves' are few sicj. supposition White 97 of pathoo _j. interactions~ of an easi)! Most! caves resui ack of ma haviors M t 968). Oth! injuries cad' anter puncf.sprained o~ keeping u cows usu, :. - - "f"" ''---~-...,....

17 ia into!-.: h~n\ ~. : behn' tr 'waiving ; of high ibou are og 968:. ~ 976). ition can s (Skoog and the maturity '. t popua red, t is juce and 'uations ack the g during niparous nen.t for :d either ier and at east and rear >u poput forage \'e a true caribou. >rtage of ed birth owever, ~ack of ave not.e to aj produt;-- dent. t caribou n eves troing ortaity. >w evef fecttve. pitation in the fonn of snow cover. icing. freezhig t;:jns. and cod rains than by a other environmert t fact;rs. To some degree. snow and ice detennine v, o.;er;:. r C W. and if caribou can ive for most of the year. P~ ~cipitation that greaty restricts forage avaiabiity combined with mv temperatures and high winds is most ikey to resut in high mortaity. Proonged and severe restrictions of forage suppies ead to manutrition and subsequent death by starvation or predation. Athough there is itte evidence for starvation caused by adverse weather conditions m free-ranging caribou on tht! mainand, there is evidence for it among Peary caribou on the High A'ctic isands (Mier et al a). beieve. as did Skoog { 968), that even if mortaity is attributed to starvation caused indirecty by weather conditions, the reguating factor is the existing weather, not the absoute food suppy. MORTALTY Caf Mortaity. High caf mortaity on caving grounds is common in caribou and is caused mosty by adverse weather and predation (Pruitt 96: Kesa 968: Skoog 968; Bergerud 97a; Parker 972a; Mier and Broughton 974). Precipitation, cod. and wind are a deady combination for newborn caves. often resuting in hypothennia (Banfied 954a) or respiratory probems such as pneumonia (Mier and Broughton 974), and subsequent death. Soft, deep snow traps caves and t}.,.e_y subsequenty die {Kesa 968). Weather aso severey imits the avaiabiity of forage to parturient cows during gestation and indirecty causes caf mortaity (Mier et a. a). This condition is criticay important to Peary caribou in the Canadian High Arctic (Thomas et a. 976, : Thomas and Broughton 978). Predation on some caving grounds can be severe and often exceeds the woves' needs (Mier and Broughton 974). WoJves. bears, ynxes, and eages seek caves as easy prey and often have marked impacts on caf crops. Predation on newborn caves varies from popuation to popuation and within popuations from year to year. t has been suggested that woves ki mosty those young that woud not survive anyway. Athough woves woud readiy take sick and ame caves, they must take heathy caves when there are few sick or ame animas avaiabe. An interesting supposition proposed by K. A. Neiand (Kein and White 978) is that woves perpetuate the occurrence of pathoogica agents in caribou popuations by their interactions with caribou and, thus, assure themseves of an easiy obtaim:"'t food source. Most accidenta deaths and some predation of caves resut from the caf's investigative behavior, its ack of materna care, or the agonistic or escape behaviors of other animas (intraspecific factors, Skoog 968). Other deaths are due to drownings, tramping injuries caused by hostie aduts (foreeg kicks and ;r anter punctures), fas from ciffs, and injuries such as.sprained or broken imbs that prevent the caves from keeping up with their group's movements. Materna cows usuay remain with their injured caves, but such CARBOU (Rangifer tarandus) 937 infants usuay fa easy prey to woves. bears. and!>ometimes eages. Poor nutrition in mothers can resut in the death of the mother before parturition. mapresentation. stin birth. -'hysioogica disorders of various organ system!;. physica abnormaities of the caf. or neonata death through manutrition of the fetu~. Mothers can transmit fata diseases t'j their young. The frequency of disease in caves is not known. but epidemic bruceosi> has occurred in barren-ground caribou of the arctic herds in Aaska (Neiand et a. 968). Adut Mortaity. Factors that predispose caribou to death by accident or predation are!heir investigative behavior. their nutritiona state. adverse weatherprecipitation. cod, and wind-predation by woves and bears. and imited forage. Forage is imited possiby by production, but usuay by avaiabiity truy the resut of weather. Much of the adut mortaity due to accidents and predation is the direct or indirect resut of intraspecific behaviora and physioogica characteristics. nvestigative behavior can ead to drowning by faing through weak ice, fas from ciffs, and snow sides. Sprained or broken imbs from trave on dangerous terrain eads to subsequent death by predation. Agonistic and reproduction behaviors may resut in encounters that ead to death by kicking or anter punctures, or cause injuries that ead to subsequent death by predation. Exampes of mortaity proximay caused by wof predation but utimatey the resut of intraspecific physioogica characteristics incude:. as bus buid up fat ayers before the rut, they can run ony with great difficuty and tire quicky, making them more vunerabe to predation 2. prime bus are often very ean after the rut and they do not buid up sufficient reserves to carry them through to spring: thus. they are more vunerabe to predation 3. near-term cows suffer from the arge size of the fetus and high energy requirements in ate winter: thus, they are more vunerabe to predation (Skoog 968). Environmenta (interspecific, Skoog 968) mortaity factors shoud take the greatest to on caribou numbers over the ong tenn. There is a great dea of debate over what are the principa mortaity factors operating in caribou popuations. Sometimes the same bioogists advanced different reasons for the various herds. Many eary workers (Leopod and Daring 953; Edwards 954: Scatter 964, 967a) beieved that range deterioration, caused mosty by fires and and cearing and ogging, on southern ranges was the principa cause for the decine of caribou numbers throughout much of North America, Cringan ( 957) and Banfied and Tener (958) bamed much of the decine on overhunting arrange detenoration. Banfied (954a), Banfied and Tener (958), Sonnenfied ( 960), and Bergerud (974a) a beieved that predation by woves or humans was the principa cause. Ony one thing is certain: at some time or another a of these mortaity facto'~, aong with weather and disease, have had marked impacts on caribou numbers.

18 -, 938 ARTODACTYLA PREDATON. Caribou evoved with the wof. and ony in reativey recent times has the wof been extirpated or seriousy reduced in numbers on the southern and eastern ranges of caribou. Wof-densities remain high among most barren-ground caribou popuations. They are possiby high reative to the ow numbers of Peary caribou, and muskoxen are a buffer prey species in High Arctic regions. Moose (.4/ces aces). sheep ( 0\ is, and goats ( Oreamnos americanus> are buffer prey species for woves on most Aaskan and Yukon ranges. The ynx has repaced the wof as a major predator of caribou caves in Newfoundand and is possiby important on other southern ranges.. The grizzy bear is an important predator on caves, and occasionay aduts, on some ranges in Aaska. the Yukon Territory. and the western District of Mackenzie in the Northwest Territories. This chapter is concerned with the management and conservation of caribou as a renewabe resource, not merey with caribou as an entity in arctic and subarctic natura systems. Bioogists are interested in the sustained use of the resource by humans as we as the continued perpetuation of the species (which incudes wof predation on caribou). Therefbre. the concept of dynamic equiibrium is not a functiona basis on which to accept the wof-caribou reationship. suggest that woves shoud be a part of the caribou's environment. But woves are arge c...-:tivores that require considerabe amounts of meat and protein for their reproductive success and surviva. A wof requires -4 caribou annuay (Carke 940; Kesa 968; Parker 972b), if caribou are its major prey. Food habits of woves on barren-ground caribou range in Canada have been reported on hy Kuyt (972). Large numbers of woves have a potentiay arge impact on a caribou popuation. based on estimates of a wof's required annua ki of caribou and the fact that woves sometimes ki more than they need, especiay of caribou caves. Less than wof per 00 caribou coud contro the growth of a barren-ground caribou popuhttion (Parker 972a ). Skoog (968) stated that predation was the greatest singe mortaity factor in most wid popuations of Rangifer. Athough Criser ( 956) and Kesai ( 968) stated that caribou, except the incapacitated and caves, coud normay outrun singe woves, Murie (944), Banfied (954a), Skoog (968), and other fied workers observed that woves are capabe of overtaking heathy caribou. Such feats are not aways necessary, as woves often take prey by surprise or ambush. When in pursuit, woves aso take advantage of any tactica mistakes the caribou may make, for exampe, eaving hard snow and attempting to cross soft snow areas. The most interesting aspect of wof hunting tactics is their abiity to run reays and to invove severa members of the pack in herding and ambushing prey. Skoog ( 968) suggested that woves can obtain their prey as needed from whatever caribou are avaiabe, and that the fina seection is made more by chance than by design. Sick and ame animas may be seected, if avaiabe, but if the caribou popuation is heathy and there are few incapacitated individuas. the woves wi take heath: animas. Woves can and sometimes do contro an,: even depress caribou popuations. PATHOLOGY. The importance of disease and parasitism ur; m caribou and as a contro of caribt'h numbers is,.~t yer we known (Kesa 968: Skm';: 968). A detaied bibiography of parasites. disea!->e:-.. and disorders of Rang{fer and other unguates wa~ compied by Neiand and Dukeminier ( 97:!>. Disea:-t and parasite burdens in caribou are endemic and ot norma incidence in most popuations. Sporadic or possiby periodic outbreaks of disease of epidemic proportions probaby occur in some caribou popuations. The ony epidemic disease in a free-ranging popuation of caribou occurred in Aaska (Neiand et a. 968: Skoog 968). Brucea organisms were associated with orchitis':'epididymitis. bursitis-synovitis. and metritis, singy or in combination in Aaskan caribou (Neiand et a. 968). Examination of barrenground caribou in Canada ed to the concusion that bruceosis was not a serious threat in caribou (Broughton et a. 970). Another important dis::a.se in Aaskan caribou is necrobaciosis (Skoog 968}. The causative agent (Spherophorus necrophorus) usuay enters through esions in the feet or mouth. and resuts in hoof rot or necrotic stomatitis. Banfied (954a) considered that actinomycosis (umpy jaw) was widespread in Canadian caribou. More recent work by Mier et a. ( 975 b) and by Doerr and Dieterich (979) has suggested, however, that most, if not a, mandibuar deformities are not of actinomycotic origin, Fibropapiomas have been found in :\askan (Skoog 968) and Canadian ca.ribou rbroughton et a. 972). The caribou is hosr to reativey few species of externa parasite, but the species that do parasitize them are common and cause considerabe harassment during summer. Harassment by boodsucking and parasitic fies can ead to injuries when the caribou react by fight, and subsequenty to death by predation. Severe attacks by mosquitoes (Cuicidae) and back fies (Simuiidae) have reportedy resuted in the deaths of caribou (Skoog 968). The warbe fy ( Oedamagena tarandi) and the nose bot ( C ephenomyia trompe) parasitize caribou (Banfied 954a; Skoog 968; Kesa 975), but they are more important in infuencing the ecoogy of caribou than as pathoogic agents (see "Ecoogy" section). The effects of the warbe fies on caribou are unknown, but they ikey cause genera debiitation of the host. Heavy infestations of nose bot arvae severey distress caribou and may weaken infected animas. nterna parasites incude various cestodes, some nematodes, and one trematode (Banfied 954a; Skoog 968). The common forms are Taenia hydatigena, found mosty in the iver; T. krabbei, mosty in the musce; and Echinococcus granuosus, mosty in ungs and occasionay in the iver. Again, the importance of these interna parasites is not known but they have genera debiitating effects on the host and coud vredispose the host to predation. The thread "' ungworm debiitated Skoog heavy there is no in barren- 967 and on some o number or.. treachero ' ice, or by sheves (

19 ' CARBOU (Rangifer tarandus) 939 rdke heathv contro and, isease and of caribou! 968: Skoon ""!S. disease!.. ' guates wa' /72. Dbe.~:~e!mic and of Sporadic or Jf epidemic! bou popua, free-ran~ing fneiand e~ ns were asis-synovitis. in Aaskan n of barreneusion that in caribou ati"e agent ; through e- hoof rot or sidered that ad in Cana :mer et a. ( 97g; has mandibuar Fibropapi- ; 968) and species of : parasitize harassment Jcking and rhe caribou "predation. and back n the deaths ii) and the tze caribou 5), but they ecoogy of )]ogy., seccaribou are :>iitation of ae severey animas. odes. some d 954a; re Taenia -. krobbei, rranu/osus'.jer. Again, not known :m th~ host The thread ungworm (Dictyocauus zadweni) is associated with debiitated caribou and supposedy causes their death. Skoog { 968) suggested that this parasite might cause heavy monaity to nutritionay stressed caribou. but there is no evidence for it. Besnoitiosis has been found in barren-ground caribou in Canada (Choquette et a. 967) and Q fever in Aaskan caribou (Hopa 975). ACCDENTS. Athough accidents usuay contribute itte to the overa monaity within a caribou popuation. on some occasions accidents can resut in a substantia number of deaths. Animas may drown whie crossing ueacherous stretches of river, by waking onto weak ice, or by becoming trapped in the water by high ice sheves (Kesa 968 and others). Many accidents occur when caribou are in arge migrating aggregations and traveing over unfamiiar terrain. They often resut in injuries that increase vunerabiity to predation. PHYSOLOGCAL Physioogica disord~rs cause very few deaths among adut caribou. Most such deaths occur among femaes experiencing compications in feta deivery. Additiona mortaity can resut from organ faiures brought on by stress in caribou of either sex or any age. SEX AND AGE DETERMNATON To evauate fuy the status of any group of caribou. the sex and age of individuas must be determined by aeria and ground surveys (Davis et a. 978b). Segregations based on physica appearances of free-ranging caribou by sex and age at different seasons of the year invove a high degree of subjectivity. The observer must note: ( ) the presence or absence of anters: (2) the reative size and deveopment of anters: (3) the age of anters-new. when in vevet. and od, when hard and poished: (4) the reative body size of individuas: (5) the state of peage condition and mot. presence of white mane on adut maes in autumn: (6) the shape of the face, forehead to nose ine, roman nosec. or straight faced; (7) the presence and deveopment of the udder; (8) the presence or absence of the vuva; (9) the presence of a penis sheath; and ( 0) indirecty, the probabe sex and age casses of companion animas (other group members). Surveys are made during spring migrations. precaving, caving, postcaving. midsummer migration, prerut, rut, postrut, fa migration, and winter. Aeria surveys of and ground counts on sma and units might give criticay miseading ratios of sex and age casses in terms of the entire herd. The probem of nonrandom distribution of caribou by sex and age can be overcome through () repeated aeria surveys of arge and units and (2) ground counts of high proportions of the herds during the rut, when the sex and age casses are the most mixed. Currenty. the most accurate method for determining the age of a caribou invoves microscopic examination of histoogicay prepared sectkms of mandibuor teeth (McEwan 963; Mier 974a, 974c). Maxiary teeth coud be used but management techniques are usuay restricted to the coection of ony the de mary bone. Cementum annui are counted but resut!- can he vaidated ony by examining teeth of known-aged caribou: this has not been done as yet. More data must be obtained on the possibiity of individua variation in apposition of denta cementum and controing interna and externa factors of cementa! growth. :-.any tooth sections cannot be read without some degree of subjectivity and annuus counts may not be exact. Most age assignment!- shoud be thought of as the age. pus or minus one year. The cost. time. and eaborate equipment required to determine the age of caribou by histoogica techniques make routine use of those procedures on arge sampes. such as annua hunter kis. questionabe. A second technique-the visua, gross examination of the state of eruption and attrition of the mandibuar tooth row (Chateain 954; Watson and Keough 954; Bergerud 970: Mier 972, 974c) is a that is required for most management procedures. Visua examination can be suppemented with inear tooth measurements to hep improve the accuracy of the age determination (Mier and McCure 973: Mier 974c). The permanent mandibuar teeth erupt during the first 3 years of the caribou's ife. Most caribou have a fu set of mandibuar teeth by the 29th month of ife (see Mier 974c). Differentiation between 2 and 3 year ods can be made in the fied on the basis of ack of wear on the posterior cusp of the third moar. Aso. open apica areas occur on the roots of the fourth premoar and third moar at 2 years (versus wear and cosure at 3 years) and can be checked with ease in the fied on dried mandibes and in the aboratory on fresh mandibes. Because the abiity to evauate wear patterns varies among investigators. it woud be more reaistic to assign caribou to age casses rather than specific years. First the caribou's age shoud be estimated to the cosest year ( - 0+ years). then the individuas paced in an age cass: caf.. 2, 3, and 0+ years. t is most difficut to differentiate between 3 and 4 year ods; however, suggest that it shoud be done. to minimize the chance of umping some 2 year ods in a 3 to 5 year ods cass. n Aaska, Chateain (954) deveoped age cass groupings of L 2, 3, and 0+ years; Skoog (968) aso produced the same age casses but incuded a caf category. Miseading estimates of age based on visua examination of mandibuar teeth coud resut from ( ) maoccusion from misaignment of the maxiary and mandibuar tooth rows, (2) variations in the curvatures of the rami. (3) and variations in primary and secondary axes of the mandibuar bades. Some caribou woud show extreme variations in attrition because of missing teeth and/or abnorma aignment of the tooth rows. caused by orientation of the teeth in their sockets (Mier 974c). Sex can be determined between oder (4-5+ years) bus and simiar aged cows and a younger animas of either sex by comparing mandibe engths

20 940 ARTODACTYLA (Bergerud 964b: Mier and McCure 973: Mier 974c). The mandibuar bade is measured from the posterior rim of the ramus to the anteriormost posicion of the aveoar bone beow the first incisor. A serie~ of such measurements must be estabished for each herd studied. deay. those measurements woud be obtained from jaws coected from animas of known sex and dates of death and known age as determined by annui counts in the denta cementum. ECONOMC STATUS n the past, caribou were a-important to arctic and subarctic native cutures: caribou meat was the stape diet for both humans and dogs: caribou hide and sinew provided the materias for cothing. tents, seeping bags. etc.; and hides, sinews, anters. and bone were used for many toois and weapons. Caribou. especiay the migratory herds, were the key to the very existence of m.dt and ndian cutures on the centra tundra and adjacent taiga (Kesa968:206-26, ): ndians in the northern borea forests might have survived without the caribou, but to do so they woud have had to ater their cutures m: kedy. An abundance of caribou meant good times and a shortage meant hardships and sometimes sow death from manutrition and starvation. Caribou remain to this day a stape in the diets of many northern peope and are sti vaued as an integra pan of their chosen (traditiona) ways of ife. Unti those native peope choose an aternative ife stye that does not have a great cutura dependency on caribou, it wi be difficut to measure the repacement vaue of caribou. Current evauations of the worth of caribou must incude moraistic and esthetica considerations. The caribou's potentia to reproduce and in turn the potentia of the offspring to reproduce and expand in numbers make today's vaue of caribou ony a mere fraction of the projected vaue 00 or more years from now. and at best a poory based guess, Another nebuous costing exercise is to estimate the esthetica vaue of caribou. The price woud be appreciabe, even at today's rates. But is this marketprice approach to the vaue of caribou a egitimate consideration beyond the point of sustained annua yieds and/or recreationa! pursuits? The Garibou is a natura entity in arctic and subarctic environs. Woud those regions not ose their very character with the passing of caribou from their andscapes? MANAGEMENT AND RESEARCH (Much of the foowing text on management and research is iberay awash with my persona opinions and beiefs; the materia does not refect :he opinions, beiefs, or poicies of the Canadian Widife Se~ic~. the Department of the Environment, or the government of Canada.) Most attempts at managing big game speci~s in North America on a bioogica basis have been ~tagued by overriding sociopoitica considerations. No other unguate species has suffer~d as much from sociopoit~ ka pres~ures in the face of opposing bioogic~! concerns as the migratory caribou of Nonh America. t i:-. impossibe truy to manage caribou unti then:~ b r.:onrro of the harvests. f we r.:annot reguate the harvests of caribou on a bioogica basis. we cannot hope to manage the resource. The management of caribou in N<mh America is unique in thut. in addition ro being seasonay sought after by meat. troph)-. and white subsistence hunters. caribou remain a srape in the diets of many northern natives who have buit their cutures around the species. The native harvest of caribou in Canada is amost unrestricted. but in Aaka there is some reguation (Davis et a. 978a). The freedom to harvest an unrestricted number of caribou has proven detrimenta to caribou popuations since the introduction of the rife (e.g.. Banfied 957: Banfied and Tener 958: Kesa 968). Most native popuations are growing and are now concentrated in settements from which organized hurtts by snow machines or aircraft are increasing the ki. The shift in most native communities to a partiay wage-based economy has increased the native ownership of rifes. snow machines, and even aircraft to be used in hunting. n particuar. aircraft hunting eads to excessive harvests of caribou because of the range that can be covered and the ease and speed with which carcasses can be returned to the settement. Native peope often state their desire to continue their traditiona ways of ife, and few peope woud deny them that wish. Unfonunatey, however. the natives' tradition now incudes the use of pow,er boars to repace kayaks. snow machines and aircraft instead of dog teams, and high-powered rifes with teescopic scopes instead of bows and spears. Caribou popui.\- tions cannot withstand unreguated harvests empoying such modern equipment and ogistics. Native hunters are not convinced that they are misusing the resource. and instead have bamed industria exporation activities and rhe activities of caribou bioogists for decining numbers and changes in the distribution of the herds. Current sociopoitica stands by native groups over aborigina rights and and caims require pubic maintenance of this conviction. n Canada. the probems of overharvest are unikey to be resoved in the near future. At present, the ony acceptabe approach is to attempt to educate native peopes on the abiity of.caribou popuations to maintain their numbers at various eves of surviva and mortaity. Natives must understand that caribou are a future resource that can be maintained ony if repacement equas or exceeds osses. The diemma is whether the migratory herds of caribou can survive unti the natives accept the truth! A second, somewhat unique consideration in the management of migratory caribou is that many migrations take caribou across nationa and internationa boundaries. For exampe, () barren-ground caribou of the Porcupine herd (00,000) move among Aaska, Yukon, and Northwest Territories; (2) barren-ground caribou of the Bevery (94,000) and Kaminuriak (38,000) herds move among the Northwest Territories, Manitoba, and Saskatchewan and in some years north- neg bioi< '.

21 ' CARBOU (Rangifer tarandus) 94 gica connenca. umi ther.: :guiate tht we cannot gement of ~- in adc.iat. trophy. :in a St'pe have buit han e~;r of in Aaska 78a). The of caribou ions since 957: Banrive popu ;ed in setby snow he shift in age-based of rifes, j in huntexcessive at can be carcasses continue >e woud ever. the wer boats ft instead eescopic popuanpoying they are!d indusf caribou.!s in the u stands d caims tion. n ey to be v acceptpeopes ain their Ortaity. Jture reacement!ther the! natives n in he any minationa ribou of Aa~ka, -ground inuriak ritorjes, s northeastern Aberta; and (3) woodand caribou on the George River herd ( 85.OOO move between Quebec and Labrador. Remnants of the Sekirk Mountains herd (20-30 sti move among southern British Coumbia. northern daho. and northwestern Washington. Those transboundary movements nece!>sirate agreement and cooperation at a eves of government and between the various widife and other resource agencies and the peope concerned with the wefare of caribou. their habitat. and the peope that use those resources. Webaanced prog:-nms are difficut. if not impossibe. to achieve when confronted with a mutitude of socia, economic. and poitica concerns among the different jurisdictions. Management Regions. One of the first probems faced by managers of caribou is to determine what unit of caribou of what range used by caribou represents a manageabe entity. Fortunatey. because of their gregarious nature and genera affinities for migration paths and caving grounds, most groupings of woodand caribou and barren-ground caribou are discrete. - enough to be recognized by managers as herds or popuations. Unfortunatey, caribou bioogists are not aways in agreement as to what constitutes a herd or a popuation. Athough definitions do exist. they are not expicit enough, without considerabe quaifiction. to justify a strong stand on the matter. n Aaska, bioogists have beieved that movements among their caribou units are frequent and great enough to cassify them as herds, and they think of a earibou in Aaska as a popuation. n Canada, bioogists working on barren-ground caribou first t,hought of them as herds; then after earning more. they thought of them as popuations (infrequent and negigibe movements among units). Now some bioogists in Canada have reverted to the herd category, based on no apparent bioogica reasoning. but just a matter of preference in definitions or iterary stye. The distributions of caribou can be considered on a geographica rather than a subspecific basis. athough distributions of subspecies co, for the most part, parae geographica distributions. suggest three management regions for caribou. First are the arctic isands cf th~ Canadian archipeago, where a rigorous poar desert,environment with unique cimatic conditions periodicay reduces surviva of Peary caribou. Those stringent cimatic factors maintain numbers of Peary caribou beow theoretica carrying capacities by causing considerabe variation in forage avaiabiity from year to year. Restrictive snow and ice conditions make estimates of carrying capacities based on standing crops and annua productions of itte or no vaue. t is ikey that the key to springtime.surviva for Peary caribou is the amount of bare, windbown beach ridges and sopes that is avaiabe to them, though the vegetation on such sites is reativey sparse. Peary caribou have been thought of as isand popuations. But recenty, Mier et a. ( a, b) and Mier and Gunn ( 978) have given evidence for extensive interisand movements (migrations) and have suggested thar Peary caribou have actuay estabished interisand popuarions. Second. the mainand tundra and taiga sections of the open borea forestc; of Aaska and Canada form a region for barren-ground caribou and migratory popuations of woodand caribou. Environmenta and socia pressures are common throughout that vast area of North America. and managers seek answers to basicay the same questions whether the caribou are in Aaska or Canada. Migratory caribou form arge aggregations that create such specia range probems as tramping or possibe oca overgrazing or continued access to ranges. Third is the. management region for the more sedentary woodand caribou of the borea forests, The geographica distribution:: of woodand caribou. their differences in foraging habits.!htir sociaization, and their year-round habitation of forests set them apart from popuations of barren-ground and woodand caribou which inhabit the tundra during a or part of the year. The genera ack of arge socia aggregations and a greater dependency on arborea ichens for winter forage are characteristic of forest woodand caribou. suggest that this subdivision into three management regions is a desirabe preiminary step for managers of caribou in their thmking and panning of proper management procedures and goas. Most management probems w~t.hin each of those areas woud be simiar enough to warrant exchange of information and ideas by a North American managers of caribou popuations. Bioogists must not ose sight, however, of the important differences between each area, such as: () seasona quaity of forage; (2) seasona avaiabiity of forage; (3) eves of harassing insects; (4) importance of cimate; (5) variation in factors causing or potentiay causing range deterioration; (6) kinds and numbers of anima predators: (7) eves of hunting pressures: and (8) amount of industria and resource deveopment. Land areas within states, provinces. and territories are most often divided into widife management regions, zones, areas, or units with itte or no ecoogica consideration given to the divisions. Usuay such jurisdictions exist mainy for the supposed expedition of administrative and manageria concerns. Therefore, whie the bookkeeping of renewabe resources may be based on such units of and, about the ony management poicies that may refect bioogica considerations are the seasons and bag imits. n reaity, the bioogica reason(s) for estabished seasons and bags may extend we beyond the boundaries of a jursidictiona unit but is usuay confined to the unit for ease of appication and enforcement. Other bioogica considerations do not necessariy reate to the jurisdictiona boundaries but are actuay tied to different ranges and management regions as have previousy defined them. Management Practices.. Two important conditions are necessary for the proper management of caribou: () the authority (abiity) to reguate th~ harvests of a. t..., - ~ ~ ~- - - c " -- ~---;-"-., -!'

22 ~:' ;"":-:: :~-:.: :\~,~...: ;..,;~~~:~...;!, '*- ' -~,.....,,... :~- ~..:::" _-:G..,-; >i '!: = -. ''.! f,.! ' i! 942 ARTODACTYLA ~ users. and (2> a bioogica basis upon which to set the reguations. For the most pan. neither has been obt:.incd~ unti the harvest of caribou by native peopes is reguated. there can be no true management of caribou as a renewabe resource on a sustained yied basis. This beief was strongy put forth for barren-ground caribou in Canada by Banfied ( 957> and Kesa ( 968)-and apparenty fe on deaf ears. as no concerted actions have been taken during the ast 20 years. However. we must continue to work at obtaining a better bioogica basis from which managers can draw. when and if they are given the opportunity to manage. n Canada, education of natives to the reaities of current caribou ;tiization wi be a ong-term process. t is un!ikey. if the educationa p;ocess is successfu, that!~ wi be accompished without some c-a.."ibou popuations first being amost extinguished. Evett a catastrophic oss of caribou woud not necessariy convince the natives of the need for reguation of harvests, for bec<!use of th~;ir beiefs, they most ikey woud not recognize or accept the cause of the oss. Therefore, educationa programs. out of necessity, maj become the major concern of caribou managers in the 980s. Management of caribou. ike that of other game species. has generay foowed a five-step sequence (Kesa 968): () reguation of hunting~ (2) instituting predator contro; (3) giving specia status to and areas for widife; (4) transpanting and reintroductions: and (5) pacing contros or: ~;e environment. REGULATON OF HUNTNG. This step has been ony partiay impemented. as native peopes are not subject to such reguations. at east in Canada. The situation appears better in Aaska. but it is sti far from being totay satisfactory. Kesa ( 968) noted that most caribou hunters traditionay, and in disregard for the aw. have habituay kied c.ribou in excess and often wasted their kis. Enforcement of existing reguations on caribou hunting has been. with few exceptions. inadequate or nonexistent. n Aaska prior to 925 itte contro was exerted on the kiing of caribou. Passage of the Aaska dame Law in 925 created the Aaska Game Commission. Subsequent reguations were basicay restrictive in permitting human harvest of c~bou unti 959. Statehood in 959 brought in argey defensibe reaxation of reguations in view of an apparent increase in the caribou popuation from 947 to 960. The ibera seasons and bags remained in effect unti it was. too ate for the Fortymie caribou herd in 973, the NechinJ! herd in 972, and the western arctic herd in That is, human harvests exceeded the annua increments in those herds, either aone or in conjunction with wof predation (J. L. Davis persona communication). Harvests were monitored in some areas south of the Yukon River as eary as 963 by mandatory harvest report cards, hunter check stations~ and viage harvest ogs. The same systems of monitoring harvests are sti in effect, with the addition of some pexmit hunts. After 975 harvest permits or harvest repons were aso required north of the Yukon River. except by residents of management units numbers 25 and 26. n Canada some contro of hunting on woodand caribou has been in effect througho Jt ra~ges during the mid-900s. Most province~ and territories cosed or drasticay shonened their seasons and reduced their bag imits in or afier the 950s. Most of those restrictions were ony paniay effective. at best. as they actuay restricted ony the nonresident hunters. Loca residents. Metis and natives. continued their traditiona hunting practices in disregard of the reguations. Peary caribou have been reativey free of hunting pressures unti recenty. soey because of their remoteness from human popuation centers. except for two nuk se!tements. Residents other than nuk in the Canadian High Arctic are subject to seasons and bag imits on Peary caribou. But in reaity there is vinuay no enforcement. The exception woud probaby be in the two settements. mainy be-cause everyone s awareness of each other's activities woud dampen such iega taking of caribou. Barren-ground caribou in Canada were hunted without constraint unti the 950s. Restrictive measures in the fomt of seasons and bag imits have affected mainy nonresident hunters and resident nonnative hunters. the atter to a esser degree, Enforcement of the restricti~ns in a manner that woud contro residents, especiay Metis. actuay is acking, with few exceptions. Treaty ndians in Canada remain above a aws and reguations p~rtaining to seasons and bags for caribou! Athough the nuk do not have a treaty, they are accorded the sau!e rights where hunting is concerned. n the 960s, as a resut of the caribou crisis. the barren-ground caribou was designated an endangered species under the Nonhwest Territories Act. Caribou hunting was restricted by aw to ony those peope that dt:. ended on them for subsistence. Reguations were passed against the wasting and abandoning of caribou and against the feeding of caribou to dogs. Widife officers tried to persuade natives not to ki cows and caves. The Canadian Deprtment of ndian and Northem Affairs during the 960s encouraged natives toreduce their take of caribou by providing them with fish nets and with commercia meats. n the 970s much of the good was undone by the Department of ndian and Northern ArTairs and the government of the Northwest Territories as they shifted their emphases to participation in utiization of caribou through organized hunts. Athough the barren-ground caribou remained on the endangered species ist, its meat was egay offet'ed for sae in the Northwest Territories. Caribou management programs in Caii~!i@ through the 960s and the 970s, for the most part, acked direction and common goas. The attitude that widife had to pay for itsef became prevaent in many agencies, and the caribou popuations have borne the brunt of this approach supposedy to justify their guardianship. beieve that this poor-man farming attitude is detrimenta to the we-being of the resource and is essentiay a vioation of the caribou manager's responsibiity to his charge. The manager's roe is twofod: essentiay he shoud manage for ( ) annua sustained yieds of the resource for those that need it and for those that de arge herds, :uion and. source. The and enjoyme One ve constraints is range. Again matter. n ony areas and Game tion. Now. Endangered jurisdiction Aaska. n Department cunai nati Department support has Basic of the ment goa:.

23 CARBOU ( Rangifer tarandus) 943 g on woodianj,\ng!:. during the ' vnc:-. do'>ed \)f ;j reduced the;r ',')f those resrnc ':!!>\. a~ the~.~~: ' : hunters. Lo.:a r d'. tnetr rra mona! 'guations. J t're.-: of huming ' se of ther rt! :er!. except ior han nuk in the. easons and ba!! ~here is vinua;, probaby be in nonna. Enforcement dd contro re:-iking. with fev.!main above a ns and bags for e a w-eaty. they mming is conan endangered ~ Act. Caribou ose peope that guations were ing of caribou dogs. Widife ki C:.QW!- and!an and North J natives to ret hem with fish undone by the,ffairs and the as they shifted tion of caribou barren-ground pecies ist, its 'orthwest Terns in Canada the most part, e attitude that vaent in many ave borne the ify their guardnine attitude is!source - and is ager's respon )e is twofod: nua ~ustained eed it and for those th:u desire to use it. incuding maintenance of arge herds for nonconsump!!ve users: and (!) conser YUton :.0d. \Vhen necr-,sary. preservation of the resource. The common e:&a shoud be the maximum use and enjoyment of the resource. not rura enterprise! One very important factor in the setting of harvest constraints is contro of the and over whic~ the caribou range. Again. Aaska has aso had the advantage in this matter. n the past. U.S. Park Service ands were the onv area:-. over which the Aaska Department of Fish and Game did no! have widife management jurisdiction. Now. however, with the 02, BLM Organic Act, Endangered Species Act, etc., the matter of who has jurisdiction over widife management is in question in Aaska. n Canada m0st ranges of barren-ground caribou are controed not by widife agencies but by the D~partment of ndian and Northern Affairs. That the best.interest of the caribou is being served is questionabe, because the department is aso responsibe for the wefare of the native peopes of Canaca and for contro of resource deveopment of the North. As a consequence. neither the N.W.T. government nor the Department of the Environment can act effectivey to curtai native hunting without fu support from the Departmem of ndian and Notthern Affairs-which $Upport ha!' not been offered. Basic to a caribou management pans is contro of the resource. Without such contro, any management program wi fa short of its utimate desired goa: rationa. sustained use of the resource and its conservation. Aso of prime importance in the management of caribou is the prote :tion of the arge herds and their migratory habits. Unfortunatey.,Jrocedures for obtaining these goas extend far beyond bioogica considerations. Socia and poitica pressures at au eves are acting against a successfu outcome of the probem of contro of the resource. Somehow. the natives must be convinced that - 2ame management a!encies are acting on their behaf '-' that suggested courses of action to manage caribou are necessary. so that their great gnmdchidren wi at east have caribou around them and be abe to hunt them, even if they probaby wi! not be abe to subsist off them on a year-round basis. At the same time. caribou must be conserved as a natura entity in their environment. This wi be by no means an easy task in view of the current resource deveopment on some of their ranges. PREDATOR CcNnroL. Predator contro in the past has essentiay meant contro of woves (the currenty favored expression is.. maintenance of the predator popuation"). Wof contro on caribou ranges has foowed the usua sequence of bounty payments, systematic government contro through empoyment of predator contro officers; the dropping of bounty payments (Pimott 96; Kesa 968), and then no contro. Bounty payments are at best a form of rura wefare; they have never proven effective as a means by which predators can be controed. t is aso ikey that no good professiona bounty hunter ever put himsef out of business! n Aaska. the U.S. Predator and Rodent Contro Program began an active campmgn of wof contro on Aaskan caribou ranges in 947. The predator contro work (i.e.. wof contro) incuded poboning. shooting from airpanes. and the payment of bountie~. t was continued throughout most caribou ranges unti 960. When Aaska gained statehood in J59 there was a dramatic shift in management emphasis. n genera. wof contro ceased. but some taking of woves \Vas sti aowed by issue of aeria wof permits. Aeria hunting of woves was curtaied nonh of the Brook!, Range in 970 and statewide in 97'2. Locaized, imited, and cosey reguated aeria wof contro has been pursued since 976 t0 hep ;::ertain depressed moose and caribou copuations to iw::r~use. n Canada, government contro of woves began in the eary 950s. when the decine of barr~n-ground caribou appeared to be catastrophic. n western Canada a serious and widespread rabies epidemic gave it a big boost in 957 (see Pimott 96 and Kesa 968 for more detais). Poisoned baits (akaoida strychnine and sometimes the poison 080) were used amost excusivey in the provinces and the Northwest Territories from 95 to the mid-960s. The Canadian Widife Service ran the contro program in the Northwest Territories from 956 to 959. then it was taken over by the Northwest Territories Northern Administration Branch. The main thrust of the contro program took pace in Manitoba and the Northwest Territories from 955 to 960. The wof contro program was considered successfu in giving the barren-ground caribou a reprieve (Pimott 96 and Kesa 968). The degree of success of.the program. however. coud not be measured quantitativey: there was a known take of 6,890 woves in the Northwest Territories aone between 952 and 96. Wof contro on caribou ranges in Canada during the ate 960s and throughout the 970s has been unsystematic. oca. and sporadic. Wof bounties have been reinstated. dropped. and reinstated soey as a resut of sociopoitica pressures. with no rea concern for the bioogica impications of such actions. The sustained annua ki of woves by nuk hunters in the Northwest Territories gives evidence that the b~unty system is not an effective way of reducing popuations of woves. As stated by Pimott (96). the bounty system does not actuay contro woves. it does not concentrate effort where it is reay needed, and animas are kied in vast areas where no kiing is justified. beieve that a web thought out and propery executed wof maintenance program is a justifiabe management too, when annua net osses of a caribou popuation constanty exceed annua recruitment. Such a program woud have ong-term benefits for both the caribou and the wc!ves. Pimott (96: 50) summed up the matter nicr.y when he said, "as ong as contro of tundra woves is judicious it wi have itte or no effect on their status dec:;.des hence. Their numbers are so cosey reated to those of the barren-ground caribou that their utimate fate hinges more on the caribou than on contro. f the caribou survive and continue their f-. '![

24 ' 944 ARTODACTYLA = migrations. the rundra woves wi survive. For this rea!on. intensive. short-term contro of tundra wove!:> cun conceivuby benefit the woves themseve~. Thi!>!iame view was deveoped and hed by the members of the Aaska Conservation Society (Weeden 976 when they caed for a carefuy executed wof reduction program on the range of the western arctic caribou herd by the Aaska Department of Fish and Game in 975. From a widife management standpoint it is unfortunate that wof contro as a management too has become such a misunderstood and distastefu exercise to usuay we-meaning but often misinformed and misguided individuas. Not ony the ay pubic but aso many widife bioogists have apparenty ost their objectivity in the :na~:er. Emotions run high. from moraistic stands to rhetoric taught by bioogists about how woves ki ony sick. od. and some young prey. There is no quicker way to cause a ook of shocked disbeief than to te a defender of the widerness that woves quite often ki heathy adut caribou. Disbeief wi ikey turn to horror if you add that the kiing by those woves often appears unnecessary, as the woves sometimes feed itte or not at a on the ki. t wi be a ong road back for both ayman and bioogist. But think that the sooner we view the matters of predation and contros in a reaistic manner, the sooner we wi reaize that both woves and caribou benefit from the maintenance of wof numbers. This consideration must, of course. be quaified to point out that wof maintenance is oniy desirabe when and where caribou popuations are in steady decine caused or acceerated by wof predation or hunting by humans. Of course. constraints on the harvests by humans in such situations are aso a necessary and vaid management procedure. n reaity, there are ony two factors in the ecoogy of caribou that we can actuay contro when given the proper authority-woves and humans (hunters, nonconsumptive recreationaists, deveopers. and expoiters). LAND AREAS WTH SPECAL STATUS FOR WLDLFE. Caribou have benefited, at east indirecty. from the estabishment of specia widife management areas, sanctuaries, and preserves. One of the best exampes is the Arctic Nationa Widife Range: 3.6 miion hectares in northeastern Aaska, set aside in 960. Caving grounds and summer ranges of the Porcupine caribou herd occ!jr within the Arctic Nationa Widife Range. But Prudhoe Bay ies ony about 00 km to the west of the refuge, and oimen beieve they can sme oi underneath it. So the future of t}:- Arctic Nationa Widife Range as a widife refuge is doubtfu in the face of strong pressures by the petroeum industry to expore and deveop any reserves within the refuge. n 926 the Arctic sands Game Reserve was created. t covered a of the Peary caribou ranges on the arctic isands of Canada and some barren~ground caribou ranges on a arge section of centra mainand tundra in the Northwest Territories. Natives coud hunt within it but no others. However, it was rescinded in 966. The Theon Game Sanctuary wa~ estabiished in 927 in the centra mainand barren-ground~ of the ~orthwest Territorie~. ts primary purpose wa!> t prutect muskoxen. but barren-ground caribou of the Bevery popuation summer and sometin~e~ cave within the sanctuary (Hoare 930: Carke 9--0J. Current~. the minera industry i!- obbying to rescind the Thehm Game Sanctuary so that they can expore it and deveop any worthwhie finds. unfortunatey, ~uch arcus actuay offer itte year-round benefit to caribou popuations: J the caribou use them ony seasonay. then range over unprotected areas; (2) natives are not restricted fron; hunting caribou on those areas: und (3) most of the estabished refuges and sanctuaries are in areas that were not heaviy hunted by natives. ~igratory caribou cannot be protected effectivey by reserves. except seasonay and ocay, because of the huge areas over which they range (Kesa 968). However, protection of caving grounds, postcaving areas, and migration routes by specia reserve and status or by specia and use reguations coud be beneficia in giving materna cows and newborn caves an added degree of protection during those time periods {Mier 974b). Such areas were traditionay afforded protection by their remuteness and ruggedness, but the airpane and snowmobie now make them readiy avaiabe. Encroachment by resource expoiters and the added pressures of a ooming North American energy crisis make the specia status of widife areas tenuous at best. f we ose the arge migratory herds of barrenground caribou in Canada and Aaska. the arguments for keeping out or restricting expoitation and deveopment activities on tundra ranges wi be greaty weakened. t is indeed a vicious circe: the natives are jeopardizing the herds by overuse, but their dependence on caribou gives the species high poitica profie that they woud not otherwise have. TRANSPLANTS AND RENTRODUCTONS. As Kesa ( 968) mentioned. to hep assure suc.~;ess wien transpanting barren-ground caribou. the restocking shoud be done in an area where () the caribou cannot stray; (2) thrre is itte or no harvesting; and (3) there is a nonconficting need for caribou. Few such paces exist for barren-ground caribou in Canada. Bergerud ( 974d) noted that, in the reintroductions of 226 woodand caribou to 8 sites in Newfoundand between 96 and 965, the greatest probem was preventing adut caribou from straying. He observed that handreared caves did not attempt to migrate or stray from the transpant sites. Bergerud (978) reported that success had varied among the transpants and to date caribou had been harvested from four transpant sites; he suggested that reeases shoud succeed on isands free of disease and predators (Bergerud 978). Caribou caves were introduced to Adak sand in the Aeutians during 958 and 959. Bergerud (978) reported that harvests have been a& high as 30 percent on the Adak popuation in 973 and that the 973 harvest was a record for sustained yied. This assume!i thatthe Adak popuation maintained its size after 973. r anj ~ ~:: a"und tay: m.: n: fj~\f;; km :- them a.j~yl c.trib they therr tent it b~ io are n is ju~ deer ieve carit icti< Co:--: VrO supr be i of c acti tive - -- ~ ; ,... ~---- ' :! :{

25 ''! '!j ",. 'i '!!!' m t<.h~j of rhe '.. : p..(\... the Be- :! within m-entt}. Tht:ton Jeveor ~;- iine 'J th~.. ver un- : m hunt- '! estab- "ere not nnotbe dyand :h they :aving fes ny e reguwsand during "j, wer~ d i ~teness t! n~>w (.. Jd the j pnergy : nuous ~arren / ments d de!reatv es ar~ iiepenti prot!isa i tran:. houd ~tray: ;! is a exist erud oadween nting andfrom suedate ;ites: a.nds td in 978)!< cent 973 mes 73. Kesa ( 968) pointed out that Southampton sand in northwestern Hud~on )> Bay '"'a!. an exceent ~ite ior the reintroduction of caribou. Caribou were abundant on the isand unti 924 but were amost totay gone by 930. The Canadian Widife Service at the request of the government of the Northwest Territories captured 52 caribou on Coats band (about 80 km south of Southampton sand) and transpanted them to Southampton sand in summer 967. No adequate!\urvey has been done of the reintroduced caribou to date. but sporadic observations suggest that they are apparenty doing we on Southampton. Restocking of isands and possiby sma areas of the mainand with caribou may have some imited potentia as a management practice. The main aim shoud be to prevent oc:aoss of caribou so that such practices ar:! no: necessary. f restocking or creating new herds is justifiabe, we shoud remember that domestic reindeer are not a egitimate substitute for caribou. beieve that the introduction of reindeer on former caribou range by any widife agency is indeed a dereiction of their charge and an irrationa act. CONTROLS ON THE ENVRONMENT. Contros of environmenta factors that adversey affect forage suppies or promote disease or parasites may at times be important and even necessary for sustained harvests of caribou. Of even more importance is the contro of activities by a users, consumptive and nonconsumptive, and especiay of industria uses of caribou ranges. Most attempts at any physica contro of the environment shoud be made ony during the most intensive management programs. with the probabe exception of forest fire contro. Forest fire suppression on winter rang~s of caribou wi be necessary if the rates of widfires increase over past rates and the areas burned are excessive in size. Kesa (968) beieved that the need to suppress fires on caribou winter ranges was web recognized, but opposition to that beief has been raised by Bcrgerud (97a, 972, 974a, 978) and others. Kesa's ( 968) pan caed for mapping unburned winter ranges and ranking them in reation to () quantity and quaity of ichens present; (2) their ocation reative to caribou use; (3) potentia effects of fires on each mature forest area; and (4) reative potentia of fire-damaged immature forest. As yet, the topic of the impact of forest fires on caribou winter range remains open to debate. Forested winter ranges coud be fertiized, but the vaue of such action and the possibe undesirabe side effects are as yet unknown. Such extreme management practices demand study on sma areas before extensive programs are considered. Fencing of hazardous water crossings has been suggested by many interested groups. Such ventures woud be very costy and probaby woud create as many hazards as they removed. Kesa ( 968) beieved that the ong-tenn average of caribou ost annuay by drowning did not exceed the number taken by ony a few native hunters. He further pointed out that caribou aso drown on quiet water crossings when struck by sudden gaes or by breaking through unsafe ice (Kesa968). Diversion of caribou CA&BOU (Rangifer tarandus) 945 fro:n one area may simpy cause their deaths esewhere, At the current and foreseeabe future eveb of caribou management Jt is hard to conceive of extensive use of environmenta contros in caribou management. n fact. it is difficut to >Ce any. unes)> rapid industria deveopment bring!> network!> of roads and pipcinej> onto caribou ranges. Aterations wi be made then ony if the funds come from mdusrry. n such an event. it may be necessary to buid crossing devtces to aow the free fow of caribou past roads and pipeines. n that event. we wi find out if caribou can Jive with modem man and his activities. think that proper management of caribou shoud incude () contro (reguations) of a persons utiizing or in some way affecting the resource~ (2) periodic. accurate estimates of the number of births and deaths in the popuation; (3) periodic, accurate estimates of the size. sex, and age structures of the popuation; (4) sampes of physica condition, when and where possibe. to gain insight into caribou performances under various environmenta conditions and to detect stress situations, when they occur: and (5) ongoing studies to detect herd spintering or dispersas that ead to egress or ingress. To date. such a we-rounded program on a periodic basis has not been maintained for a the major caribou herds or popuations. Probems inherent in most fied work are ack of funds, Jack of continuity in personne and programs, shortage of experienced personne, and insufficient preparatory training in procedures for fied techniques. Many fied techniques demand powers of observation. Most observers can improve their skis with practice and proper training, but some peope are incapabe of becoming accurate observers regardess of their forma training. speciaized training. or time in the fied. Practica training courses and fied supervision often ead to the detection of pe0pe who are potentiay good observers. and those who are.. bad" can be assigned other tasks. Many, if not most, previous estimates of the different parameters for measuring and evauating caribou popuation dynamics have suffered from ~wo basic shortcomings: () the sampe sizes reative to the size of the popuation were too sma; and (2) sampes were often ocaized and represented ony a singe or, at best, a few segments out of a the segments that made up the popuation. Caribou are usuay strongy segregated and, think, probaby remain segregated to some degree. at virtuay a times. Therefore, sampes that are taken from ony one or a few restricted areas are not going to refect accuratey the condition being measured. This has been we documented in the centra arctic caribou herd of Aiaska (Cameron and Whitten 979; Cameron et a. 979). Future work must stress the need for arger sampes and better representation of the popuation in samping procedures. suggest that any sampe that is ess than 0 percent of the popuation size and not taken from, at east, most segments of the popuation be used with extreme caution; arger, seemingy mote representative sampes shoud sti be -

26 946 ARTODACTYLA suspect when setting management poicies pertaining to han em regimes. The caribou manager shoud never torget that he i:. working ony with estimates that are subject to wide variances, usuay of unknown magnitudes. Bergerud ( 978). among others. has stated that census resuts shoud not be accepted without two independent methods showing agreement. basicay agree with this beief but at the same time reaize that it is ften difficut to obtain one good estimate under fied conditions. There is itte wisdom in reducing the effort in one sampe just to take a second sampe. When funds and manpower prohibit two samping procedures at high eves of effort, i think it woud be better to go for one sampe at a. ~igh eve of accuracy than two at reativey ow evds. Whie the use of two independent methods is a!ways desirabe, many management decisions wi often have to be made on the basis of ony one such estimate. Therefore. fied workers must strive for continued improvements in methodoogy. DETERMNNG SEX AND AGE STRUCTURE OF THE PoP ULATON. The best time to perform ground cassification of caribcm for sex and age composition is in the autumn. usuay ate October through to mid November. The actua degree of segregation of the caribou at that time is unknown. Many workers beieve that the caribou are not segregated near the beginning of the rut. t is unikey that this assumption is true, as have found that groups of caribou kied during prerut were sti highy segregated in ate September (Mier 974c). t is probaby true. however, that there is ess segregation at that time than at other times of the year. Sex and age cassification can aso be done aong with caf production counts (from the ground) just after caving. The resuts can be compared to autumn compositions for further insight into evauations of estimates of sex and age composition of the popuation. Caribou are cassified as buis, cows, yearings. and caves, An unknown category is usuay necessary if you are going to account for a animas examined, f time and experience permit, you shoud separate bus into mature, prime bus (fu-sized bodies and anters; often roman-nosed head profies) and young bus (smaer bodies, with sma anters; Jacking face profies of oder bus). Some attempt shoud be made at identifying juvenie maes (one to three years od), which may be confused with adut cows. You can ony be sure of adut cows that have been sexed by externa genitaia. dentification of sex and age casses shoud be based on externa body characteristics, especiay externa genitaia, as outined by Bergerud ( 96, 964a), Skoog ( 968), and others. Experience, if not essefttia, is very vauabe, and the same peope shoud do these counts each year. Bergerud (974c) suggested that the sex ratio of one mature bu per two mature cows is a species characteristic for caribou. Thus, he suggested that sex ratio shoud be sought if the quaity of the stock is a primary consideration in the management program. Popuauons of caribou that are heaviy sport hunted i(r trophy unimais are not ikey to maintain this sex rati of one marure mae per two femaes unes.!> conservative seasons and bugs are imposed. Sex and age compositions of native kis are most ikey as much by chance as by design. A vuiabiiry to the natives usua) dictates what is taken each year. Natives did and prohnb!y woud sti foow the fat cyce in rhe caribou. if possibe: that is. they woud seect for mature maes in summer unti the rut. then they woud seect for mature cows throughout the winter unti just before parturition. t is obvious that there was no room for any concept of conservation as we understand it in the native's use of caribou on an annua basis. The kiing of pregnant cows throughout the winter was reasonabe when the natives had to ive by the od rue of "a bite of fat for each bite of ean." This rue was based on sound judgment. acquired over many years of coping with the extreme cod of the arctic and subarctic winters-intake of fats were mandatory to surviva. Unfortunatey, this practice of kiing co\\ s throughout the winter persists in the absence of a tru~ need-od habits die sowy. nformation on sex and age of.humer ds coud be used to gain some insight into the compo~ition of the popuation. However, the accuracy of such information must aways be questioned and treated with caution. Data on sex and age composition can be obtained by setting up hunter check stations on roads that give imited access to areas where arge sport or nonnative subsistence kis of caribou occur. n the past and currenty, coecting of sex and age statistics from native kis of caribou was usuay not even attempted. This was the case for good reason. as most often the obtainment of figures for the annua harvest of caribou by natives was done ony once a year. Such exercises reied on the natives' having good memories and a wiingness to be cooperative and accurat~ in their accounts. Such reiance was sedom fruitfu. More recenty. some attempts have b!en made at coecting ki statistics severa times a year or even monthy. Future efforts at obtaining statistics on the native harvest of caribou shoud incude sex and age information, when possibe. However. we must first get accurate figures of the annua harvest of caribou by nativ~s before we give much concern to the sex and age of their kis. This seemingy can best be done by empoying some respected native resident of each settement to make weeky door-to-door checks of the hunters' success. t is unreasonabe to expect most native hunters to provide accurate information on sex and age compositions of their annua ki of caribou if they are asked ony once each year. When and if such information is obtained, the bioogist must remember that the sex and age compositions are measures of the ki and not necessariy of the popuation as a whoe. The constant segregation of caribou makes it highy unikey that the kis wi represent the entire popuation. An exception to this condition may occur when many entire socia groups a e kied seasonay. Some bioogists subscribe to.the use of reativey sma shot sampes of caribou to evauate popuation que~tion or wrong \ popuati unfortunat sampes h But that i inherent i i popuatio Bani buit ife Banfied mandibe mortait fied's ( 99 perc When ( 955) tion of shot sa ponder The

27 ffi! h\!.. ~! ) ' t 'Pi. if! ~ tn ur;: uriany na ; of Lbe on ing eric ~~a. ~~ ud ~.of d f?r- 'tth ob :hat naage not. as ua e a 0d acom.ue ven the age 'irst by age :mtemt :ive age are the po-. of of. ep -'0- are use ;ate popuation dynamics. A:-; mentioned previousy. yu~stion sut:h a practice, even though have done it out of nece~suy. \Ve reay never knov. when we are right or wrong unti we have samped most or ai of the popumion under consideration. However. accept the unonunute fact that we often have to work with sma sampes because they are the best that we can obtain. Bm that is no reason for osing sight of the dangers inherent in such use. especiay when appied at the popuation eve. Banfied (955) and Mier (974c> have both buit ife tabes for barren-ground caribou in Canada. Banfied constructed a time-specific ife tabe from 292 mandibes found in 948 and 949 on summer and winter ranges of caribou. used 943 caribou kihed between March 966 and Juy 968 (Mier 974c). A of my sampes were taken from the Kaminuriak popuation on both winter and summer ranges. Banfied determined the ages of his specimens based essentiay on the pattern of wear and eruption known for white-taied deer at that time. sexed a of my specimens in the fied. did histoogica sections of mandibuar teeth to determine their ages by counting annuations in the denta cementum. then determined eruption and wear patterns for the mandibuar teeth on a seasona basis. Banfied ( 955) presented ony one curve for both sexes, whereas my sampe aowed the construction of a curve for each sex (Mier 974c). Tota monaity was 92 percent for the first 0 years in Banfied's ( 955) ife tabe and 92 percent for femaes and 99 percent for maes in my ife tabe (Mier 974c). When think about the apparent quaiity of Banfied's ( 955) sampe of caribou ower jaws used in construction of his ife tabe and the supposed quaity of the shot sampe of caribou that used in my ife tabe. ponder whether shoud concude that the two :,ampes were equay good or equay bad. Athough the resutant information, for the most part. ooks reasonabe, there is no way of knowing whether or not it represents the iving popuation. Therefore. even though such exercises are interesting to do and tak about. they must be treated as suspec! with regard to whether or not they truy refect what they are supposed to represent. EVALUATNG PHYSCAL CONDTON Whenever the opportunity presents itsef, bioogists shoud necropsy caribou and record infonnation on physica condition, cause(s) of death, and occurrence of disease and parasitism. Bergerud (978) suggested that animas shoud be coected in May and June to evauate the annua nutritiona impact of overwintering and again in Sept~mber for statistics on conditions deveoped in summer. agree that such sma sampes provide the bioogist with additiona feeing for his work and may refect conditions being experien.;!ed at the popuation eve. do not share Bergerud 's opinion, however, that such sampes refect conditions appicabe to the entire popuation. My ack of faith in such sma sampes is especiay strong when the information is used for evauating popuation dynamics. The recent histories of a animas in a caribou popua- CARBOU (Rangifer caru;;.d-u3) 947 tion are not the same: ( a caribou in a popuation do not winter or summer on the same areas. so they do not necessariy obtain the same quaity and quantity of forage: (::!) they are not necessariy exposed to the same eves of disease and parasitism: 3J a caribou in a popuation are not exposed io the same eve of predator and insect harassment: and (4 caribou are usuay segregated by sex and age. Therefore. infonnation from a sma sampe. especiay from a singe segment of the popuation. may be quite miseading. Granted. if unfavorabe conditions are detected in a sma sampe. then greater efforts can be made to determine occurrence throughout the popuation. For these reasons suggest that bioogists may justifiaby use sma sampes for gaining in.sight into subject areas of concern or possiby ;n the hope of detecting unfavorabe conditions that are suspected because of the prevaiing environmenta conditions. But if managers use the information from such sampes as the soe means of making management decisions, they run the risk of being seriousy wrong. Caribou bioogists and managers must remember that they are working with a iving entity and thus some, possiby many, detrimenta impacts brought on by unsound judgments in management decisions may be irreversibe. Sponsoring organized hunts in Canada for the soe purpose of obtaining caribou for natives that had no caribou readiy avaiabe to them tends to be a bemish on our records as professionas. think this because those hunts were carried out with no knowedge of and probaby no rea thought for the abiity of the caribou being harvested to sustain the additiona ki~ no bioogica information was obtained from the kis~ and on some occasions, the carcasses or portions of the meat were eft behind and never subsequenty retrieved. Organized hunts are by no means traditiona or esthetic and are offered to the natives as privieged services. not as a fufiment of their native rights. Therefore, such hunts shoud be aowed ony when the widife agency concerned knows that the caribou popuation receiving the additiona harvest can sustain it. The hunts shoud be controed by widife personne in a manner that aows the coection of as much bioogica information as possibe. When a widife agency (or any other government agency) provides such an expensive service, maximum benefit to caribou bioogists as we as the natives shoud be the outcome of the venture. LVE-CAPTURE AND MARKNG OF CARBOU Many projects require ive-capture and marking of or teemetric attachments on individua caribou. Such activities are often the most difficut and sometimes the most expensive phase of a project, and they are aways integra to the success or faiure of the study. Longand short-term studies of the discreteness of herds, subpopuations, and popuations for specific ranges and caving grounds; dispersas of individuas or groups from their major socia units; seasona movements (rates of trave) and distributions; group cohesion; companion animas; causes of natura

28 948 ARTODACTYLA i t.... mortaity: hunting pressures: reproductive success of tndividua femaes-a these studies require individua!~ identi iabe caribou tor group identification and ottt!n teiemetricay equipped caribou for detaied. usefu resuts. Capture and markjng techniques are in a constant state of deveopment. as is teemetry. Capture techniques for caribou can be considered under two major headings: ( ) water cros~ing, and (2 on and. Lh e-capture at Water Crossings. Captur-e is greaty faciitated if the caribou under study have known. traditiona paces of water crossings on rivers or akes that are wide enough (usuay 400 m or more) to aow pursuit and overtaking of the caribou whie they are swimming. The basic technique that has proven successfu for capture of caribou at water crossings is as foows. First. the capture team must know approximatey when, where. and in which direction the caribou wi be crossing the water body. Stabe boats, 7-m freighter canoes with 20-hp outboard motors or the equivaent. are positioned severa hundred meters (or coser if cover is afforded) from one or both sides of the anding site on the far shore of the crossing. The capture crew remains out of sight. or at east inactive and on a vantage point, if one exists. When the caribou have entered the water, the crew waits unti the caribou have swum we out into the crossing. Once the caribou are far enough from shore that they can be overtaken before they can retreat. from the water or make it across and out of the water, the crew(s) take to their boat(s). The idea team size is three four-person crews each consisting of one motorman. one recorder, one croaker and tagger (the person who initiay catches and subsequenty ear tags or affixes the coar, etc. to the caribou), and one hoder (the person who takes the caribou from the crook and hods it in pace whie it is being worked on) in three boats. One boat races to the far side of the crossing from where the caribou entered the water to prevent a re~ treat: the other boats fank the swimming caribou. f necessary, one of the fanking boats races ahead to tum the caribou back if they are getting too cose to their anding. Each motorman singes out an individua caribou, pus aong side of it, and throttes back to the swimming speed of the caribou, The croaker reaches out with his shepherd's staff and crooks the caribou around the neck and pus it to the gunwae at the bow of the boat. (A asso can be used instead of a shepherd's crook.) The hoder reaches over the gunwae and grabs the caribou by the head, ifts the caribou sighty, and hods it securey against the top of the gunwae. The tagger frst reaches over the. side and grabs the caribou's tai with one hand and the outward hind eg with the other hand. He then ros the caribou up and toward the boat to expose its ana region and genitaia, and then cas out the se;~c to the recorder. Both the tagger and the hoder determine the age cass of the anima and ca it out to the recorder. The tagger then.reeases the rear of the anima and moves forward to attach the eanags, coar, etc. The caribou is then reeased and the motorman pus the boat away and pursues another anima. or maneuvers to prevent animab from eaving the water. There <:an be many vuriatiom. on this theme. f manpower is in short :>uppy. a three-man crew \Vithout a separate recorder can be used. Even a two-man crew coud do the joh on wide crossings: one motorman t with a speci:j! device for ocking the boat on course after the caribou is caught). who moves forward to do the tagging. and a croaker who aso serves as hoder and possiby recorder. f there i~ a need for capturing and hoding animas for subsequent work or transport. corras can be buit out of sight on the far shore and boom-type fences extended out on the water. The caribou coud then be herded inside the booms and run into the corra. Entire groups coud be captured if there was need for such a arge-scae operation. Fixed-wing airpanes and heicopters on foats may be used when the capture of ony a few caribou is desired and mobiity is a primary consideration. When using an aircraft, the piot ands and taxies up behind the caribou so that the anima comes between the foats. The w.;;ge" works from one of the foats or on some fixed wing 3:rpanes from the cross-arm attachment betwet!n foa~): (Staying out of the prop is mandatory for the success of the operation). Often the anima can be worked on without being restrained, but, if necessary. a asso rope can be dropped over its head and the anima secured in pace by it. There are many advantages to capturing caribou at water crossings: ) a arge number of caribou can be captured within a reativey short period of time; (2) the technique is apparendy ess stressfu to caribou than other capture techniques (minima restraining time, no drugs are used. and individuas are aowed to remain with or catch up with other group members immediatey after reease); (3) manpower can be as ow as two men or as arge as resources permit; (4) the cost per anima caught is usuay much ower than by other techniques: and ( 5) the technique can be made fairy mobie. especiay if the capture of a arge number of caribou at each ocation is 0t a goa. The main disadvantage of reiance of capture of caribou at water crossings is that the technique is restricted to use during the open water period, which is usuay ony from Juy into September of most years. Aso, the caribou to be studied must use suitabe water crossings with reasonabe annua reguarity. There is one sociopoitica disadvantage to a methods of ive-capture and marking or radioequipping of caribou: most northern natives object strongy to such management procedures. Athough most; if not a, northern natives beieve that the kiing of caribou is a nobe activity, they aso, unfortunatey, beieve that the marking and reease of caribou for whatever purpose is degrading to those caribou, Many northern natives hod a mystica beief that marked caribou wi te others where they were debased and thus the herds wi not return to those ocations. With a strand of ogic, they concude that the kiing of caribou near traditiona water crossings is arightbecause dead caribou te no taes. The free intermixing of marked or radio-equipped caribou sho ~. \.houd argue~ individuab. origina ca~ movements anv more a:- ce~sfu wof necessary w Jea~t in Can. Caribo~' at water cro.. as both wo ' water crossi~ A tech~r: caribou into some object in!! two tog br'z>wn with attach pairs front end of, detergent ju woud then in patterns The idea em corraing c. manner that caribou. f o aso serve a( caribou wo~ they though! Live-Captu! of Uve-capn The threem: (2) corras. straint. DisL possibe tee are beyond consider or~ successfu t greatest pro NETS AND ( ered togeth overapping studies that a seasona vantage of use are usu The u~ tange nets. on the grou poes or tre from cannc ras can be ogs, using Hoding pe canvas, or ines or a cause a drc Set ta tionary mo water eros caribou wt

29 CARBOU (Rangifer tarandus). preyt:nt ani- :hi th~m.!. t, crew without! \Vo-man ere\\! n.! motorrnan om on cour\t: t'otwurd to do : 'V.!!\ a~ hoder hodinu ani ; ;orras c-an be. m-tvpe fences ' coud then be i corra. Entire ' ed for such a i ers on foats 'ew caribou is, :ration. When. ies up behind between the.! foats or on s-ann attach opj!- manda C en the anima tined, but. if over its head :ng caribou at uibou can be ftime; (2) the caribou than ing time. no ed to remain embers im- be as ow as. (4) the cost.hun by other made fairy :e number of : main disadtt water crosse during the om Juy into U to be studh reasonabe ntage to a t or radiottives object s. Athough at the kiing fonunatey, caribou for ribou. Many that~ marked debased and ions. With a! kiing of is arightio-equipped caribou shorty after reease with uncaptured caribou shoud argue against any degradation of the handed individua!.. Abo, recapture of marked caribou ar their origina capture ocation during the same seasona movements indicates that the experience does not have any more asting d'fec~ on them than woud an unsuccesfu wof attack. Convincing the n~tives of this is a necessary tasj..: f we w~mt to continue such work (at east in Canada). Caribou nonnay exhibit caution and nervousnes! at water crossings, which is ikey predator orientated, as both woves and bears often frequent the shores of water crossing areas. A technique we worth investigation is to decoy caribou into the water. have thought about taking some object such as a 0-gaon fue drum and attaching two together end to end. woud paint them dark brown with a nong3'e duck boat type of paint and attach pairs of caribou ~nters of various sizes to the front end of each pair at drums. Painted empty pastic detergent jugs coud be used as heads. The decoys woud then be anchored in ~he water off the anding site in patterns simuating groups of swimming caribou. The idea coud be appied to the use of boom fences for corraing caribou by attaching individua decoys in a manner that creates a physica barrier to the swimming canbou. f the decoys were effective, they woud ikey aso serve as a 'psychoogica barrier;'' as swimming caribou woud be ess ikey to fight the boom fences if they thought that the decoys were rea caribou. Live-Capture on Land. Tested and potentia methods of ive-capture of caribou on and are many and varied. The three main categories that come to mind are ( ) nets. (2) corras, and (3) immobiization by chemica restraint. Discussions of a of the variations of a of the pos~ibe techniques under each of the above categories are beyond the scope of this chapter. wi. therefore, consider ony those that know have been proven successfu or that think are most ikey to have the greatest promise. NETS AND CoRRALS. Nets and corras wi be considered together because many of their potentia uses are overapping. Corras are better suited for ong-term studies that require the ive-capture of many caribou on a seasona or annua basis. Nets have the primary advantage of being portabe and the associated costs of use are usuay much owe!.". The use of nets incudes set tange nets, propeed tange nets, nets fired from cannons mounted obiquey on the ground or mounted obiquey or horizontay on poes or trees at varying heights, and tange nets fired from cannons mounted on pursuing heicopters. Corras can be buit out of secured nets, wire fencing, or ogs, using meta or wooden posts or trees for uprights. Hoding pens in corras coud be covered with burap, canvas, or even pywood. Corras can be buit with set ines or automatic eyes that when triggered woud cause a drop gate or swinging gate to cose and ock. Set tange nets, tange nets propeed from stationary mounts, and corras can be used aong trais, at water crossings that are too narrow for capture of caribou whie they are sti in the water, or in "bind spots" on terrain where caribou can be herded. Tange net!. fired from cannons mounted on heicopter<; can be Ust!d anywhere and at any time that u heicopter can operate. Such caprures woud, however, most ikey b.! ess traumatic to the captured animas in deep snow. where the impact of netting woud be buffered to!-orne extent by the snow cover. Fixed-wing airpanes. heicopters. nowmobies, hoats. and peope on horseback or afoot coud be used to herd and harass the caribou into the net!-> or Cmas. Most netting operations using set tange nets woud probaby be more successfu during periods of deep snow cover. but they can be used ~t a times of the year. Cannon netting and corras can be used yearround with good success. Corras, extensive drift fences. and mazes buit of chopped and uprooted trees and bouders were constructed by aborigina natives to direct caribou to peope ying in wait with spears and bows and arrows. Snares were aso set aong the courses of the fence ines and in mazes to catch and hod caribou for kiing. The effectiveness of such structures aong migrationa paths was most ikey proportiona to the ength of the drift fences and the numbers of physica mazes and hunters. The e;dstence of such traps argues we for the ong-tetn fideity of caribou to traditiona migrationa paths. t aso suggests that automatic tagging or coaring devices (snares) coud be used with some success aong forest trais or aong trais in deep snow. Set tange nets have been used successfuy to ive-capture caribou {Des Meues ~ Mier et a. 97). The approach is best used during winter or during the eary period of spring migrations when trais in deep snow are reguary used by cariuou. The best situations for sets are on forested river or ake shores where caribou are coming onto the frozen water bodies aong we-defined trais in the snow. Likey sites for setting nets are ocated by aeria search (or can be done by snowmobie treks). Cessna 85s are we suited for the work, but other ight. fixed-wing airpanes can be used. Use of a heicopter woud give an additiona degree of fexibiity to the operation. Heicopters woud be mandatory on terrain that acked arge bodies of frozen water for anding fixed-wing aircraft. Fixed-wing airpanes on big whees can, however, be used on High Arctic isands and on some mainand tundra habitat types that are suitabe for big whee andings. Snowmobies can be used to transport netting crews and to chase caribou into the nets instead of using aircraft, especiay if the crew were to work one genera area for an extended time period. When a ikey netting site is ocated. the network of trais is then examined and a pace is chosen where a major trai eaves the forest and comes onto the frozen water surface. The pane is then anded near that point, making sure not to taxi over the major trai, and the crew depanes with the net(s). The net (a good size is 3-4 m by m with about 25-cm 2 mesh) is set across the trai just inside the trees from the shoreine. The upper edge of the net is hung on cut-off imbs and tied every 3-6 m at a height of about 2 m to secure it. The oose bottom of the net is then pued (pursed) t

30 ' -! ii - "" "" ~ ~ ~ ~- -~ ~--~----- ir,. - ~Lj 950 ARTODACTYLA. toward the direction from which the caribou wi enter, so that their feet wi be entanged withiq the mesh when their body strikes the net and causes it to move backward. Antered individuas are more apt to be caught than anieress ones. The use of a second net set 3-4-m behind the firm wi increase the catch of anteress Cand antered) animas. The effectiveness of set design varies directy with the configuration of the net cfi!wre 47.7A. Use of doube nets with a narrow entra"fice that coud be bocked off with one end of a net (figure 47.7B woud increase the catch per set and thu!> reduce the cost per anima caught consideraby. especiay if arge catches were desired. When the net is in pace, two trees shoud be cut and paced in the snow about m out on the ice to guide the piot(s) in driving the caribou into the net(s). The piot then take the pane aoft, reocates the caribou as quicky as possibe, and herds them toward the net. f the caribou have eft the ice during the time required for setting the net(s), the piot dives the pane in front of them to tum them back to the ice. Once the caribou are back to the ice, or if they have remained there, the piot ands the pane on the far side of the ---- ~-... ~ / 3 ' / './ ' \.,~ / ' / ' \ \ \ ~.,---..., ' ' / \ \ 7 \ \ \ \ \ \ \ \ \ \ \ \ / \ \,,,, \\ \\ \,, -; \\ ~,'),~.,., ' A 8, , / / 5 ' \,, / ' \ \ '\ /, , / 6 ' \ \ \ ' \ \ \ \ \ \ \., / / '' // 8 \\ \ \ \\ \\, ',, / \' \ \._,....J..,_ ~;--7'... FGURE Tange net set design. A, design used for Jive capture of caribou (Rangijer rarandus) (Mier et al 97):, ine: 2, crescent; 3, ange; 4, horseshoe: 5, obovate; 6, obanceoate. B, designs thought to he more effective but untested: 7, doube obanceoate; 8, doube obovate. caribou reative to the nettmg site and begins driviag the caribou toward the net by taxiing after them. The piot shoud hod down the taxiing speeds to aow the caribou to get onto their main back trai and foow it into the net. f the caribou attempt to eave the main trai and break for the ice on side trais, the piot shoud cut them off and herd them back to the main trai. When within m of the net, or when the shoreine is coming up, the piot shoud race the engine to induce the caribou to make a fina rush into the net. Caribou appear to see the net just before they hit it and if they are not encouraged to jump into it, they sometimes stop and wak or run around it. During herding, the netting crew remains hidden as cose as possibe to the entrance of the set without being detected (often 20m or ess, depending on cover afforded). Crew members on snowshoes cover any side trais cose by the set that caribou might attempt to use if they break from the main trai as they approach the shoreine. f caribou do break onto those side trais, the snowshoers rush out toward the caribou and frighten them back onto the main trai. Once the caribou are entanged in the net, the crew members rush in and hog-tie each anima by the two hind egs and one foreeg. The restrained animas are then processed as quicky as possibe, hed, and reeased as neary as possibe at the same time to reduce the chances of group break-up due to the capture and restraint. A specia effort shoud be made to see that a caves are reeased with their mothers, if known, or at east with an adut cow. The capture of caribou when using set tange nets shoud be about proportiona to ( ) the size, number, and configuration of nets used: (2) the amount of manpower empoyed; and (3) the number of aircraft or snowmobies used for herding the caribou into the net(s). Recent and prevaiing weather conditions. especiay the character of the snow, are important to netting success, as are piot abiities. Once temperature fuctuations cause snow to thaw by day and refreeze at night, frozen river and ake surfaces become too hard packed for airpanes to and and pursue the caribou. Aso, once the bearing surface of the snow wi! support running caribou, they can freey eave their trais and the operation becomes more difficut. At those times heicopters or snowmobies can be used for herding the caribou. A Cessna 85 or simiar airpane can carry up to a three-man netting crew, one net (3 X 30m size cass), associated equipment and suppies, pus one piot. As itte as one man and a hepfu piot can do the job, but. with much more handing time and much ess efficiency. The idea team size when using fixed-wing airpanes is three panes with tht'ee three-man netting crews: two panes to serve as "fankers" and one on drag to hep keep the caribou from breaking from or turning back on their direction of travel A Be-206B heicopter or its equivaent can carry a two-man crew and necessary materias, pus piot. When using a heicopter; manpower is necessariy reduced, but a singe heicopter with a skiied piot can perform neary as we ts tr caribou Ti e U' tange tets : ~ the cat ture proach!!\ hr as doe the i advam age C' partie~. two periods to areas such gratio.a p about on sn supp:es or. when: disn consi :ierati and au t sed~ es am The u~ shoud be mari zes fip. and decoy nati ies usc satnater i They aso gruts oft ikt bus. mcving at ' cases car (P75a) ti st)rofoam. repaceab ey.~s. and Th ~ref ore ;. \ sma canr and proce;,, t\. ".,.-' CHEMCA if tee mtque; ani nas h.: unc: er dif:: \ imnobiiz~ Hruthoom (HErthood M9t (Ho 95), su 90: Jac Hatringto~ fox and t9e5)-hi t is ~, deui her tain a ct~ Amt rican~ ( 9~ 8) a:j Mcretrid. usee for safe han. mmobii tran~port erati;ms. taini g i

31 ,, s drivino 0 em. The!.iow the foow it f! trai and 0ud cut Li!. When preine is '~o induce i Caribou td if they imes stop s hidden j t without on cover ;: any side :pt to use roach the, trais, the frighten net, the ; a ~y the ei animas and ted, e to rei e c<fpture je to see >thers, if nge nets number, t of man~ ;rcraft or into the nditions, tortant to perarure :freeze at too hard caribou. support rais and >se times rding the 'Y up to a!e cass), piot. As job, but ess effi (ed-wing n netting j one on froq or e-206b tan crew using a :. but a m neary as we as three fixed-wing airpanes when herding the caribou. he use of drop tange nets and cannon-fired tange nets has not yer been deveoped specificay for the capture of caribou. think that both netting approaches hod promise for the ive-capture of caribou. as does the Jse of auromatic tagging devices. The main advantage of these techniques woud be that sma fied parties. two-man crews or arger. coud be eft for ong periods to work caribou intensivey within specific areas such as minera icks. winter trais, and migrationa paths. n restricted areas, crews can move about on snowshoes and hand pu their equipment and suppies on sedges. When working in arger areas where disturbance of the caribou is not as much of a consideration, the crews coud trave by snowmobies and hau their camps. equipment, and suppies by sedges attached to their snowmobies. The use of baits and decoys for attracting caribou shoud be investigated. Kesa ( 968:2 i 5-26) summarizes findings of severa authors on the use of baits and decoys by primitive northern natives. Primitive natives used sprinkings of dog urine, vegetation. and satwater ice bocks to attract caribou to pitfa traps. They aso decoyed caribou in summer by imitating the grunts of cow-caf pairs and during rut by beowing ike bus. Caribou can even be attracted to a person(s) moving about in the open: waving white coth often causes caribou to come in cose to investigate. Bubenik (975a) used dummies of caribou heads made of styrofoam or PVC-foam pruty covered with nyon fur, repaceabe anters of various sizes, and movabe ears, eyes. and neck to approach caribou in his studies. Therefore, it shoud be possibe for bioogists with sma cannon nets and a ot of patience to deve;bp sets and procedures for attracting caribou to the sets. CHEMCAL MMOBLZATON OF CARBOU. No other technique for ive-capture of free-ranging or captive animas has been as widey used by as many bioogists under differing conditio.s and on different species as immobiization by chemica restraint. Books (e.g., Harthoorn 970, 976), a Widife Society monograph (Harthoom 965), and hundreds of artices-e.g., on M99 (Houston 970; Presne et a. 973; Coggins 975), succinychoine chor!de (Mier 968; Aen 970; Jacobsen et a. 976), rompun (Bauditz 972; Harrington 974), phencycidine hydrochoride (Stefox and Robertson 976), and nicotine (Behrend 965)-have been written on the subject. t is obvious that the subject cannot be covered in detai here. recommend that interested bioogists obtain a copy of Chemica mmobiization of North American Game Mammas, by Hebert and McFetridge ( 978) as a basic guide to the subject. Hebert and McFetridge ( 978) provide information on ( ) drugs used for immobiization; (2) modes of deivery and safe handing of drugs; (3) reactions and sensitivity to immobiization; (4) care of animas dur.. ng capture, transport, and reease procedures; and (5) ega considerations. They give 86 tabes from the iterature containing information on dosage rates, induction time, CARBOU (Rangifer tarandus) 95 immobiization time. recovery time. etc.: 2 tabes for deer. for moose. 9 for ek. 6 for sheep. for goat:.. for pronghorn anteope. 4 for bears. 2 for woves. and 6 for native feids. Their work contains ists of 29 different drugs that are sod under 45 different trade names and addresses of 28 suppiers. They aso cite over 50 references that reate to chemica restraint of mammas and provide a gossary of terms. pus appendixes of other vauabe information on the subject. Many bioogists have c:ptured caribou with different drugs in projectie syringes CCap-Chur or Pneudart trajectories) fired from C0 2 - or powder-operated rifes and pistos. Some have used syringes (darts) attached to the tips of arrows propeed by ongbows or attached to bots shot by crossbows. However. itte has been pubished in readiy avaiabe journas on the resuts of immobiization by chemica restraint of caribou. assume that this ack of iterature reating to the immobiization of caribou is due for the most part to the fact that other wokers found, as did, that they coud capture caribou using the same approximate dosages of succinychoine choride as reported by Bergerud et a. ( 964). Why more recent efforts using rompun and M99 have not reached the iterature. do not know. Woodand caribou were immobiized with succinychoine choride deivered by CO~ Cap-Chur equipment in Newfoundand (Bergerud et a. 964). Free-ranging caribou required reativey heavier dosages of mg of succinychoine choride per kiogram of bo dy weight than captive caribou. which required ony mg/kg. The caribou were ocated on foot and by heicopter searches during the caving season and the rut, then staked on foot by the shooter. Femae caribou appeared to have higher toerance to the drug than maes: captive femaes mg/kg: captive maes mg/kg: free-ranging femaes mg/kg: and freeranging maes mg/kg. Aaskan bioogists have shot caribou from heicopters using a Cap-Chur 28-gauge powder dart gun and 3-cc syringes oaded with 2-20 mg of succinychoine choride (R. D. Cameron persona communication). They varied their dosages of the drug by season of the year based on estimated body weights but continued to find, as others have, that caribou exhibit considerabe individua variation in their toerance to the drug. Bioogists in Aaska have aso used mg of M99 with mg of rompun to immobiize caribou (R. D. Cameron and J L. Davis persona communication). Rompun was used in combination with M99 to reduce the excitant effects of the M99 on the drugged caribou. t ts ikey that s+ mg in combination with rompun woud be better dosages for adut mee caribou (J. L. Davis persona communication). Haigh (976) used mg/kg of rompun to immobih~e captive caribou. Many of the other immobiizing drugs (Hebert and McFetridge 978) that have not been tested on caribou or for which resuts have not been reported wi ikey be usabe. The ive-capture of caribou by immobiization with projectie syringes has the advantage that the f

32 952 ARTODACTYLA equipment is highy portabe and rhus the method is appicabe as an opportunistic technique. Concerted efforts ~.:an be made to capture caribou hy shooting with dans or the dare equipment can be reguary transported in the fied with the bioogist and used whenever an opportunity pre~en~s itsef. This technique is we suited for the recapwre of caribou for remeasurements or refiuing of teemetric packages. Bioogists shoud remember that there is amost aways some mortaity as5ociated with the use of this technique and pan accordingy. Marking Caribou. Materias and methods for marking arge unguates are constanty changing and considerations for caribou are indeed numerous. There is no practica way of covering the subject here. Bioogists must become famiiar with the possibe materias and methods for marking caribou through the iterature and discussions with other bioogists. Possibe materias range from natura materias through space-age synthetics. Easticized synthetics can be used where expansion of coars is necessary or desirabe: e.g., on rutting bus and young, fast-growing caribou. However, if coars for adut maes are attached at a singe upper point by materia such as poyethyene braided rope, easticization of the coar is not necessary. But think a coars put on young individuas shoud be easticized. Materias used in marking must be abe to withstand extreme fuctuations in temperature and varying degrees of wetness and dryness. Many of the space-age synthetics appear best suited for proonged (two to four vears) use under :uch environmenta conditions. The basic n:ethods of marking caribou are: ( ) ear tags, (2) ear stream;:rs, (3) coars, (4) cooring with dyes or paints, (5) tattooing. (6) freeze branding, and (7) branding. EAR TAGS. Many different sizes and shapes of commercia catte and sheep ear tags are avaiabe for use on caribou. Custom stamping of ettering or numbering b _ _.,e- f\ the tags by the manufacturer or retaier. Such ear tags are usuay meta or pastic, puncture types, and are attached with specia piers or by interocking parts. Most ear tags serve mainy as semipermanent markers, as they are usuay not readiy visibe. Some arger, coored pastic ear tags woud be visibe to ground observation, at east over short distances. Th~ bioogist must remember that the arge meta tags coud ead to the freezing of ear tissue at the extremey ow temperatures commony experienced by caribou in winter and thus subsequent oss of ear tags. EAR STREAMERS. Most ear streamers are made of pasticized materias but other space-age synthetics wi probaby do the job as we, as most are resistant to deterioration at ow temperatures. They can be coor coded for identification of individuas. Ear streamers can be attached by passing the streamer through a sit in the ear, then back through a sit in the upper end of the materia (see Gies 969:30, figs. 8, 20) or they can be attached with rivets or interocking or sef~ ocking fixtures. CoLL\RS. Most of the coaring materias and coar designs used on other arge Nonh American unguates t:an be adapted fri; use on carihou. The materia u~ed and the design and size Gf the coar woud be governed mainy by the use that was going to be made of it. The two basic considerations are its visua quaities and its physica properties. f the primary use of the coar is going to be for visua identification of individuas. the body of the coar shoud be wide, ettered or numbered with a penetrating paint. or coor coded. f the primary use is to be as a carrier for some kind of teemetric package, strength and position (fexibiity) mainy shoud be sought. n such cases a narrow bet type of coar may be more satisfactory. Of course, a combination of high visibiity and strength for retention of a teemetric package might be necessary or desirabe. have found viny-coated. nyon-webbed materias most suitabe for coars, but any of the space-age synthetics woud probaby do as we or better. We have successfujy used.eather beting, industria canvas beting. viny-coated nyon fabrics, neoprene-coated fabrics, and hypaon-coated fabrics. Other fabrics that can be used are neoprene-hypaon-coated nyon, coo-top marine fabric, viny/nyon aminate, riviera vinycoated dacron, fuorescent viny-coated conan, f uorescent/ uminescent viny!-coated cotton, and fuorescent viny-coated nyon. Poyethyene ropes make exceent coar attachments; they are simpy tied and the ends heated and meted back into a section of the rope above the knot. Coars can aso be affixed around the caribou's neck with snaps through grommets in the ends of the coars, or by sewing, riveting, nuts and bots, interocking or sef-ocking fixtures, stapes, and adhesives. CoLORNG. Caribou can be marked by cooring them with various dyes or paints. The dyes or paints can be appied by ( ) hand throwing, {2) being propeed from co!! and powder guns, (3) using automatic spraying devices. and (4) dropping from aircraft. How. when. where. and how many caribou you want to mark wi argey determine the kinds of dyes or paints and methods used for appiying them. There are commercia paint bas avaiabe for marking animas that are fired by Cap-Chur rifes and pistos, bursting on impact and eaving coored botches on the animas. Automatic spraying devices coud be used for marking caribou at minera icks or aong estabished trais. Hand throwing or shooting coor markers at caribou woud be practica ony when a sma sampe of marked caribou was needed. A reativey cheap and effective method of coor-marking arge, gregarious unguates has been used by Simmons ( 97) on Da sheep and by Mier et a. ( b) on Peary caribou. The method empoys the use of a dye soution hoding tank carried in a sma aircraft with a quick-reease mechanism controed by the piot-much in the fashion of a crop-dusting operation. This method of dye spraying caribou has great potentia for use in studies requiring the marking of arge numbers of animas. Studies of socia or range affinitit>s from thts aeria dyt! Man~ ong-term Bergerud pears ro of individ merey eve and those bioi

33 ,-- /.:uur!j :! -., i ~~:u ~rned. The bj b. ; Ur ~!!>.the :be red mar) netric ainy pe of 'bina- :~ of a ibe.!, most ;hetics ';cess!:ting,. brics, an be ; 'o-top. ;inv- :. ~ ono.n. ~ and ropes :.y tied ion M \ffixed grometing..tures. them ::an be i from,:aying when. >k wi 's and.ercia! fired. ct and )matic bou at i:hrow- '! prac- ' u was od of ; been. Mier npoys )t sma :ed t;,y, opera- ; great Jng of range affinities or fideities by caribou can benefit greaty from thb method of marking groups of caribou by aeria dye spraying. ~t:my-probaby most-caribou bioogist~ think of caribou as being in open socia units with no true ong-term permanencye.g.. Lent J966a: Skoog 968; Bergerud 97-c. Their subsequent concusion appears to be that as there is no rea need for recognition of individuas if socia units ack stabiity-the caribou merey recognize each other at the sex and age cas!- k\e and not as specific individuas. disagree with those bioogists on this mutter, both as to the degree of openness of caribou socia units and as to the animas abiities to recognize each other as individuas (Mier 974cL think that this method of dye spraying from aircraft woud give an exceent opportunity of testing the discreteness of wintering groups of caribou during the spring migrations. Entire groups of wintering caribou on the same range coud be dye sprayed various coors (each group a different coor). Then they coud be observed as they move into arge migratory aggregations and their subsequent basic wintering group cohesiveness determined. Therefore, wi summarize the aeria dye spraying method as have used it on Peary caribou (Mier et a. b). The dye spray tank was designed by and buit to order for Simmons (97). The tank that we used con~ tained about 200 iters () of soution. t was mounted on the foor of the airpane behind the piot's seat with ug bots. Two Heio Courier aircraft were used for the spraying. Ony the piot rode in the airpane during the dye spraying "fights (Canadian MOT reguations governing the use of the tank in ight aircraft). The piot controed the oad of dye soution with an 'a or nothing'' type of quick-reease ine. That is, whenever he pued the ine, 200 iters of soution was dumped; no partia dumping was possibe. A biodegradabe, nontoxic dye was mixed in fue drums by meting snow and heating the water to about C, then addin~ the dye and stirring it into soution. aowing the snutiun to coo. and then crdding about 20 iters of isopropy acoho. The isopropy acoho aided in the rapid absorption of the dye imo the caribou peage. The dye soution was then pumped into the hoding tank in the airpane. The piot took off, ocated a group of caribou as quicky as possibe, and made a 0-20-m atitude pass over them. The pane overshot the group of caribou and as the piot judged his ead and passed by, be pued the quick-reease ine and dumped a200 iters of dye soution. The timing of the reease of the dye soution requires experience, so some tria and ei'!or exercises are necessary, and the fying requires a skied piot. Hundre\is or even thousands of caribou can be marked by using this technique whenever caribou are in arge groups, especiay whie they are on migration. The technique coud aso be used on sma groups during p~riods when the caribou are reativey sedentary. The cost per 200-iter oad of dye soution shoud be between $00 and $50, not accounting for abor and aircraft time. Each oad cou!ci mark tens of caribou CARBOU (Rangifer tarandus) 953 under favorabe conditions. Therefore. the cost per anima marked shoud be much ower than that by any other such highy visibe marking technique currenty known. TATTOONG, FREEZE BRANDNG, A'D BRANDNG. t seems that tanooing. freeze branding. or branding woud be methods of imited use in marking caribou. These possibe methods of marking caribou shoud not. however. be totay forgotten. as there might be imited occasions when they woud be appicabe to specific situation!.. For exampe. ip tattooing gives a permanent hidden identification, if one is ever needed. Ecoogica Bioteemetry. Perh:ps no singe too has as many appications to techniques for obtaining bioogica data as teemetry. Ecoteemetry has made possibe many otherwise not practica studies and has aowed us to obtain a weath of hitherto unavaiabe bioogica information. New horizons await us with future deveopment of the art. Future deveopment in miniaturization of teemetric parts, computer anayses of teemetric data, sateite packages, decassitication of miitary technoogy, etc., wi greaty expand the fied of ecoteemetry. One version has it that the art of ecoteemetry as an appied science began in 957 with the teemetering of the temperature of an incubating penguin egg in the Antarctic (Adams 965). n ess than two decades great strides have been made in the fied of teemetry. Exampes of bioogica data that can be obtained from teemetry are many, the potentia probabiities countess: e.g. body temperature, respiration rates. respiratory tida voumes. bood pressure. heart sounds. intestina pressures, ph. insuative quaity of ft::r and feathers. ectoparasite counts, information on behavior and behaviora responses to natura and foreign stimui, navigation (homing), migration, daiy and seasona movements (rates, distances. ocations. etc.), and macrocimatic and microcimatic dat;. recommend that any bioogist not we versed in eectronics find an exp:!rt in eectronics to work with him at the start of the study. My own experiences with radiotee:netry suggest that equipment faiures ;Jre not uncommon and that correction usuay reqtc>ires the expertise of an experienced eecti'onics troube shooter. nterested bioogists can famiiarize themseves with the fied of ecoteemetry by reading such materias as "A Sped' Report on Bio-Tec:metry," Bio Science (Februar., 965), for background and more recent proceedings, such as the fire:~ (Long ) and second (Long 979) internationa conferences on widife teemetry at Laramie, Wyoming, for information on the current state of the art. CURRENT RESEARCH AND MANAGEMENT NEEDS Caribou bioogists, managers, and systems modeers from North America and Scandinavia took part in a symposiun-workshop at the University Cif Aaska. 7-9 Navembei', to review their current state of kn0wedge, -v{ewpoints, and research priorities. be-

34 . 954 ARTODACTYLA ieve that th::::>se proceedings (Kein and White 9~ 8) serve as the most compete and up-to-date thinkhg on re~~t!.rch needs for caribou. t was agreed that ',mowedge of the foowing parameters was necessary for an understanding of popuation dynamics of ca:.ibou: () popuation size. (2) popuation structure. (3) agespecific birth and death rates, ( 4) mortaity to predators. (5) other natura mortaity. (6) dispersa. (7) human harvest. (8) distribution (seasona movements). and ( 9) status of other interacting herbivores ( competitors and aternate prey of predators). Bergerud ( 978) stated that. in genera, there b reason to be hopefu for the future of caribou, as they are highy adapted and adaptabe. truy hope he is right. Currenty can see itte cause for optimism. especiay with regard to the future of migratory herds of caribou. Subsistence users want to utiiize the caribou beyond the maintenance capabiities of the herds. Expoitation of petroeum, gas, mineras, and water has the potentia for disrupting migrations and reducing avaiabiity of caribou ranges: Poiticians and pressure groups through misguided beiefs or ack of understanding or indifference are wiing either to not act at a or to take the wrong actions on behaf of the caribou. Even some widife peope suggest turning the management of caribou over to the native peopesthat indeed woud not be a panacea for the caribou. f we are to manage and conserve caribou propery. a hitherto unknown. concerted. c-out effort mus~ be made-nijw. Federa, provincia, state, and cerrito, ria agencies charged with th~ responsibiity for management and conservation of caribou and th: ir runge~ must vigorousy press for the constraints necessary to protect the caribou. hope that such an effort is in the offing-and is offered in time. To par~ phrase Harper ( 955), no other arge North American and mamma is of such primary im" portance as the caribou as a source of food to natives and nonnatives; no other c:mi~s out such extensive and spectacuar migrations~ no other can be seen in such vast herds: and no othef exhibits so cose an approach to a warden of Eden'' trustfuness in the presence of man. Thus, perhaps, no other is more worthy of being cherished and safeguarded in its natura haunts for use and enjoyment of future generations than the caribou of North America. Working- with caribou has given me the opportunity to witne~s some of the most marvenus sight'> in nature, and has reinforced in me the beief that the arctic woud indeed be an empty and without the migratory caribou. am particuary gratefu to other caribou bioogists who have t&ken the time over the years to debate matters pertaining to ecoogica ar:d. behaviora reations of caribou to their environment. offer specia thank to those coeagues who argued the "gray ar~as" of those considerations, as much of what think know about caribou bas been deveoped out of those conversations. ~ especby thankfu~ tc J. L. Davis, Aaska State Department of Fish and Game, for providing me w;th a gtt!at dea of information on caribou and their management in Aaska. thank Dr. A. Gun, Northwest Tenitorics Widitc Service: M. C. S. Kingsey. R. H. Russe, and Dr. W. E Stevens. Canadian Widife Service. for critjca\' re:uiin!! eariicr \ersions of the manuscnpt for this chapter_ - aso offer a specia thanks to G. D. Hobc.;on. din::ctor. Poar Continenta Shef Pmjec. t. Energy. Mines and Re ~ources Ca:::.td!.. for over a r.:'~cade of financia and kh!t<;tica support of my work on c::.ibiu _ Without that suppon. s'omc o the work woud never have been done, and oth.-::r -.tudje: woud have been carried O't ~t much ower Jeveh of eiton. LTERATURE CTED Adams. L Progress in ecoogica bioteemetry. Bio Science (February), pp Aen, T. J mmobiization of whiie-taied deer with succinyochoine choride and hyauronidase. J. Wid!. Manage. 34: Anderson. G.: Andersson, K. Brundin. A.; and Rappe. C Voatie compounds from the tarsa scent gand of reindeer (Rangifer taf('r. :;}, J. Chem. Eco. J L Anderson, R. C. ;97. Neuroogic disease in reindeer (Rangifer tarandus) introduced imo O::~tario. Can. J. Zoo!. 49: Banfied, A. W. F The present status of North AmeriQ c;jn ca.ribou. Trans. Nonh Am. Wid. Conf a. Preiminary investigation of the barrenground caribou. Can. Wid!. Serv. Wid!. Manage, Bu.. Ser., no. JOB. 2pp. ~ h. The roe of ice in the distribution of mammas. J. Mamm. 35: A provisiona ife tabe for the barren-ground caribou. Can. J. Zoo!. 33: ~ The pight of the barren-ground caribou. Orvx 4: A revision of the reindeer and caribou genus Rangifer. Nat. Mus. Can. BuL 77. Bio. Ser. no pp The disapp::arance of the Queen Charotte s and's caribou. Nat. Mus. Can. Bu. 85: The mammas of Canada. Univ. Toronto Press. Toronto. 438pp. Banfied, A. W. F. and Tener. J. S A preiminary study of the Ungava caribou. J. Mamma. 39: B;;;uditz, R Sedation. immobiization. and anesthesia with Rompun in ~aptive and fre:e-iving wid animas. Vet. Med. Rev. 3: Behrend, D. R Notes on fied immobiizatipn of white-taied deer Wi!,! nicotine. J. Wid!. Manage. 29:889-~~0. Beanger, L. F.; Choquette, L. P. E.; and Cousineau, J. G.! 967. Osteoysis in reindeer anters: sexua and seasona v~ations. J. Cac. Tiss. Res. : Be, K. L., and Biss. L. C.. Overwinter phenoogy of pants in a poar semi desert. Arctic 30: 8-:!. Bergerud, A. T. 96. Sex determination of caribou caves. J. Widt. Manage, 25: a. A fied method to determine annua parturition rates for Newfound~and caribou. J. Wid!. Manage. 28: b. Reationship of mandibe ength to sex in Newfcundand caribou. J. Wid!. Manage. 28: Eruption of permanent premoars and moars for Newfoundand caribou. J. Wid!. Manage. 34: t ' ( ~~~~~--~-9-7 a_. f Nev. iound; f -- J7b ' \V'.dJ. ~ar (_. caribou. W ---!97::!. \Vid. :-,.ai -.; crang{er Fied :-.iat. ---!974a ing setterr h Newfound.. 974c. Pages 394 behaviour Vo.. ' c 940pp.,. 974c.. tion. mov ' Pages 55:f behaviour! Vo. 2. n CJ40pp caribou. < -- 97f caribou D. L. Gi: anr man 494pp. Bergerud. A r moose ' equipme: Bergerud, A. reared ct Oikos :2: nrough~on. ~~. :-.er. ;f wrandu.\ Rang(/ J. Zoo!. Brough:on. E Cutaneo' caribou. Bubenik, A. ife of J. R. Lt:j eds. Pre Pap. u -- 9/ g({er, H D. R. -j deer ans Rep. nd Bunne, F.. Mier, R.;Scc repon J ) manag,, D. R. ~ deer a Rep. n Caef, G.

35 CARBOU (Rangifet tarandus) 955 : L i j.:h'f.. ; Rt.: j.. ' ~ "'~ '\..... j!.: :! i J'' J 'rt Bio- ) with \'idi. / t!. c. nd of!."'75 '- --!Ran Zoo. mcri- :4-,7- ~~:: f!.:::! J, 6(t j r:e ' r~.! J.mnto i ina!) -f>73.!.thesi' mcs.. j on of!nage. J. G Jsoia i! ogy of :aves. ma~- ruund arturianage. sex -56. moars 4;96;' a. Abundance or'rorage on the winter range of Sewfoundand carib0u. C:m. Fied ~at. 85: b. Hunting of.stag canhou n ~e\\--foundand. J. Wid!. ~ana!!e. 35: c.. The popuation dynamics of Newfounci:wd caribou. Wid!. Monogr. no pp '2.. Food habits of ~ewfoundand caribciu. J. Wid!. Mana!!e. 36: t-.ovement and rutting behavior of caribou ( Rang~(er wrandus at Mount Abert, Quebec. Can. Fied ~at. 87: a. Decine of caribou in Nonh America foowing s~uement. J. Wid!. Manage. 38: b. Reative abutidance of food in wint::i? for Newfoundiand caribou. Oikos 25: c. Rutting beh::viour of Newf:Jundand caribou. Pa!!es in V. Geist and F. W.iters. eds. The behaviour of un!!uates and its tek~h.~n to management. Vo.. UCN N~w Ser. Pub!. 24. '.. xge:;, Switierand. 940pp d. The roe of the environment in tht aggregation. movement. and distur>ance behavioth of caribou. Pages in V. Geist and F. Waters, eds. The behavkwr of unguates and its reation to management. Vo. 2. UCN New Ser. Pub!. 24. Morges, Switzerand. 940pp The reproductive season in Newfo:mdap_d caribou. Can. J. Zoo!. 53: The annua :-.nter cyce in Newfoundand caribou. Can. Fiei:! Nat. 9:0: Caribou. Pag(!s 83-0 in J. L. Schmidt and D. L. GUbert. eds. Big game of North Americ~: ecoogy and management. Stackpoe Books, Hanisburg. Pa. 494pp. Befgerud. A. T.; Butt, A.; Russe. H. L.; and Whaen. H mmobiization of Newfoundand caribou and moose with succinychoine chcri~ and Cap-Chur equipment. J, Wid. Manage. 28: Bergerud, A. T.. and Noan. i'v~- ;; 970. Food habits ofhand reared carib<.~t Rar.gifer arandus L. in Newfoundand. Oikos 2: Broughton, E.: Choquette. L. P. E.: Cousineau, J. G.: and Mier. f. L. 970 Bruceosis in reindeer, Rang({er rarandus L. and the.iigratory barran-ground caribou. Rangifer to,randus grocnandicus (L), m Canada. Can. J. 2oG ~ 02. Broughton. E., Mier, F. L.; and Choquette. L. P. E Cutaneous fibropapiomas in migratory barren-ground caripou. J. Wid. Dis. 8: Bubenik, A. B. 975a. Significance of anters in the socia ife of barren-ground caribou. Pages in J. Ft. Luick, P. C. Lent, D. R. Kein, and R. G. White, eds. Proc. st nt. Reindeer and Caribou Symp. Bio. Pap. Univ. Aaska, Spec. Rep. No.. 55 pp b. Tmtonomic vaue of anters in genus Rangijer. H. Smith. Pages 4-63 in J. R. Luick, P. C. Lcm, D. R. Kein. and R. G. White. eds. Proc. st nt. Reindeer and Caribou Symp. Bio, Pap. Univ. Aaska, Spec. Rep. no.. 55pp. Bunne, F.; Dauphine, T. C.: Hiborn, R.; Mier, D. R.; Mier, F. L.; McEwan, E. H.~ Parker, G. R.; Peterman, R.; Scotter, G. W.; and Waters, J. C Preiminary report on computer simuation of barren-ground caribou m.~agement. Pages in J. R. Luick, P. C. Lmt, D. R. Kein. and R. G. White, eds. Proc. st nt. R :indeer and Caribou Symp. Bio. Pap. Univ. Aaska, Spec, Rep. no.. 55 pp Caef, G. W Popuation status of caribou in the ~orthwest Territories. Pages ~)-6 ; D. R. Kein and R. G. White. ed!-. Parameter~; of caribou popuati0t7 ecpogy in.-\aska. Proc. Sym}:'. and Wo.rkship. f.io. Pap. L:nn. Ab~ka. Spec. Rep. no. :_\, 49pp. Caef. G. W. and Lof{ie. G. r-..l 975. :\minera ick of the barren-ground caribou J. Mamma. 56: Cameron. R. D.. ond Luick./ R Seasona change~ hn wta bc'dy water. e:ma-ceuar Ouid. and bood \'Ume m grazing reindeer. Can. J. Zuo. 50: Cameron. R. P.. \Vbite. R. G.: and Luick. J. R Th~ aq:urnu;.\ttqn of water in reindeer dunng winter. Page! J'o\ in J. R. Luick. P. C. Lent. D. R. Kein, and R. G. White. eds. Proc. st nt. Remdeer and Caribou Symp. Bio. Pap. Univ. Aaska. Spec. Rep. no.. S5pp. Cameron. R. D.. and Whitten, K. R Seasona.! movemenw and sexua s~~egation of car;bou de!ermined by aeria survey. J. Wid!. Manage. 43: Cameron. R. D.: Whitten. K. R.; Smith. W. T.! and Roby, D. D Caribou distribution and group composition associated with comtruction of the Trans-Aaskan pipeine. Can. Fied Na~. 93: Chapin, F. S. ; Van Ceve, K.; and Tieszen. L. L Seasona nutrient dynamics of tundra vegetation at Barrow, A~.aska. Arctic Apine Res. 8: Chateain, E. F Anter-jaw study. Fed. Aid Wid!. Rest. F:ogr. Rep. Proj. W-3-R. 8 vos. U.S. Fish and Wid!. Serv. Juneau. Aaska. 8:4-4. Choquette. L. P. E.; Broughton, E.: Mier, F. L,: Gibbs. H. C.: and Cousineau. J. G Besnoitiosis in barrenground c:uibou in northern Canada. Can. Vet. J. 8: Carke. C. H. D A bioogica investigation of the Theion G.ime Sanctuary. Nat. Mus. Can. Bu. 96, Bio. Ser. no pp. Coggins, V. L mmobiization of Rocky Mountain ek with M99. J. Wid. Man.ige. 39: Cringan, A. T History. food habits, and range requirements of the woodand caribou of continenta North America. Trans. North Am. Wid!. Conf. 22: Criser. L Observations of woves hunting carib0u. J. Mamma. 37: Dauphine, T. c.. Jr Bioogy of the Kaminuriak popu ation of barren-ground caribou. Part 4: Growth. reproduction. and energy reserves. Can. Wid. Serv. Rep. Ser. no. 38, 7pp. Dauphine. T. C.. Jr.. and McCure, R. L Synchronous mating in Canadian barren-ground caribou. J. Wid!. Manage. 38: Davis, J. L. ')78. History and current status of Aaska caribou herds. Pages -8 in D. R. Kein and R. G. White, eds. Parameters of ;.:aribou popuation ecoogy in A;ska. Proc. Symp. and Workshop. Bio. Pap. Univ. Aaska, Spec. Rep. no pp. Davis. J. L.: Gra}.voga. C.; Reynods. H.: and Vakenburg. P. 978a. Human utiization of the Western Arctic caribou herd. Fed. Aid Widt Rest. Proj. W-7-8 and W-7-9, Job 3.20R. Aaska Dept. Fish and Game, Juneau. 43pp. Davis, J. L.: Reynods. H. V.: Vakc:nburg. P.: and Shideer. R. T. 978b. Sex and age composition of the Porcupine caribou herd. Fed. Aid Wid. Rest. Proj. W-7-9 and W-7-0, Job 3.23R. Aaska Dept. Fish and Garm, Juneau. 23pp. Dehority, B. A. 975a. Characterization studies of rumen bacteria isoated from Aaskan reindeer (Rang[fer tarandus). Pages in J. R. Luick. P. C. Lent, D. R. Kein, and R. G. White, eds. Proc. st nt. Reindeer and ---

36 956 ARTODACTYLA Caribou Symp. Bio. Pap. Univ. Aaska. Spec. Rep. no.. 55pp. --- J975b. Rumen ciiate protozoa of Aaskan reindeer and caribou ( Rmt!{ifr:r wrandus L.. Pages in J. R. Luick. P. C. Lent. D. R. Kein, and R. G. White. eds. Proc. st nt. Reindeer and Caribou Symp. Bio. Pap. Cni'r. Aaska. Spec. Rep. no.. 55 pp. Des Meues, P Operation caribou. Les Carnets de Zooiogie 25(2): Bnnging back the caribou. Animas 002): Des Meue~. P., and Heyand. J. 969a. Contributions to the studv of the food habits of canbou. Part 2: Daiy consumption of ichens. Nat. Can. 96: b. Contributions to the study of the food habits of caribou. Part : Lichen preferences. Nat. Can. 96: de Vos. A Behavior of barren-ground caribou on their caving grounds. J. Wid!. Manage. 24: Doerr, J. G., and Dieterich. R. A Mandibuar esions in the western arctic caribou herd of Aaska. J. Wid!. Dis. 5: Dugmore, A. A. R. 93. The romance of the Newfoundand caribou. J. P. Lippincott Co., Phiadephia. 86pp. Edwards, R. Y Fire and the decine of a mountain caribou herd, J. Wid. Manage. 8: Land form and caribou distribution in Britis' Coumbia. J. Mamma. 39: Edwards. R. Y., and Ritcey. R. W Migrations of caribou in a mountainous area in Wes Gray Pare. British Coumbia. Can. Fied Nat. 73: Foods of caribou in Wes Gray Park, British Coumbia. Can. Fied Nat, 74:307. Edwards. R. Y.: Soos, J.: and Ritcey, R. W Quantitative observations on enidendric ichens used as food by caribou. Ecoogy 4: Erickson. C. A Some preiminary observations on interspecific acoustic communication of semi-domestic reindeer, with emphasis on the mother-caf reationship. Pages in J. R. Luick, P. C. Lent, D. R. Kein. and R. G. White. eds. Proc. st nt. Reindeer and Caribou Symp. Bio, Pap. Univ. Aaska, Spec. Rep. no.. 55pp. Espmark, Y. 964a. Rutting behavior in reindeer (Rangifer tarwrdus L.>. Anim. Behav. 2: b. Studies in dominance-subordination reationship in a group of semi-domestic reindeer (Rangifer tarandus L.). Anim, Behav. 2: Freddy. D. J Distribution and movements of Sekirk caribou, Can. Fied Nat. 93:7-74. Freddy, D. J., and Erickson, A. W Status of the Sekirk Mountain caribou. Pages in J. R. Luick, P. C. Lent, D. R. Kein, and R. G. White, eds. Proc. st nt. Reindeer and Caribou Symp. Bio. Pap. Univ. Aaska, Spec. Rep. no.. 55 pp. Freeman, M. M. R Assessing movement in an arctic caribou popuation. J. Environ. Manage. 3: Frick, C Homed ruminants of North America. Am. Mus. Nat. Hist. Bu pp. Gies, R. H.. Jr ed Widife managemencte~hniques. 3rd ed. rev. Wid. Soc., Washington, D.C. 623pp. Gunn. A.; Mier, F. L.; and Thomas, D. C. 9H, The ~urrent status and future of Peary caribou U<angifer ra'r.mdus pearyi) on the arctic isands of Canada. Bio. Cooserv. 9: Haigh, J. C Fentany-based mixtw:es in exotic anima neurooptanagesia. Pages in M. E. Fower, ed. Ptoc. Am. Assoc. Zoo. Vet. Congr., St. Louis. 265pp. Harper. F The barren-ground caribou of Keewatin. 'niv. Kan~a. ~us. Nat. Hist. Mbc. Puh. no. o. 63pp Harnngton. R mmobiion-rompun in deer. Vet Rec. 9-+: Han. J. S.: Heroux. 0.: Cott"', W. H.: and Mib. C. A. 96. The infuence of cimate on metaboic and themu! respnnses uf infant caribou. Cun. J. Z(0. 39:S45-S56 Harthoom. A.M. Y65. Appication of pharmaceogic:t <tnd physoogica principes of restraint of wid anirnab. Wid!. Mongr. no pp. ---!970. The fying synnge. Geoffre;. Bb. L(ndon. ::!Sipp The chemica capture of animab. Baiere Tinda. London. 46pp. Heape, W. 93!. Migration, emigration. and nomadism. W. Heffer & Sons, Cambridge, Engand. 369pp. Hebert. D. M.. and McFetridge. R. J Chemica immohiiz:uion of North American game mammas. Aberta Recreation. Parks, and Wid!.. Fish and Wid!. Div., Edmonton. 84pp. Hemming, J. E. 97. The distribution and movement patterns of caribou in Aaska. Aaska Dept. Fish and Game, Game Tech. Bu. no.. 60pp. Henshaw, J. 968a, A theory for the occurrence of anters in femaes of the genus Rangifer. J. Brit. Deer Soc. : The activities of wintering caribou in northwestern Aaska in reation to weather and snow conditions. nt. J. Biometeor. 2: Conseq!ences of trave in the rutting of reindeer and caribou (Rangifer tarandus). Anim. Behav. 8: Hoare, W. H. B Report on investigations affecting Eskimo and wid ife, District of Mackenzie, , together with genera recommendations. Dept. of nterior, Northwest Territories and Yukon Br., Ottawa. 44pp. Mimeographed Conserving Canada's musk-oxen (being an account of an investigation of Theon Game Sanctuary with a brief history of the area and an outine of known facts regarding the musk-ox). Dept. of nterior. Northwest Territories and Yukon Br., Ottawa. 53pp. Hoeman. D. R., and Luick:, J, R.. Lichen species preference by reindeer. Can. J. Zoo!. 55: 368-!369. Hopa. C. E Q Fever and Aaskan caribou. Pages in J. R. Luick. P. C. Lent, D. R. Kein, and R. G. White, eds. hoc. st nt. Reindeer and Caribou Symp. Bio. Pap. Univ. Aaska, Spec. Rep. no.. 55pp. Houston, D. B mmobiization of moose with M99 etorphine. J. Mamma. 5: rving, L, 966. Adaptation to cod. Sci. Am. 24:94-0. rving, L., and Krog, J: 955. Temperature of skin in the arctic as a reguator of heat. J. App. Physio. 7: Jacobi. A. 93. Das Render: eine zooogische Monographie der Gattung Rangifer. Akad. Verag. Leipzig. 264pp. Jacobsen, N.; Armstrong, W.; and Moen. A Seasona variation in succinychoine immobiization of captive white-taied deer. J. Wid!. Manage. 40: , Johnson, D. R Mountain caribou: threats to surviva in the Kootenay Pass region, Briush Coumbia, Northwest Sci. 50:97-0. Johnson, E. A., and Rowe, J. S Fire in the subarctic wintering ground of the Bevery caribou herd, Am. Midi. Nat. 94:i-4. Kesa, J, P Continued barren-ground caribou studies. Ca.<. Wid!. Sent. Manage. Bu. Ser.,no pp. and no.. Kein. D. deer _. and G:une Kein, D. popu: shop, Krog. J. C growi P.C. ' ~~-~--~ ~~ ~-~-----~----~~-~

37 ~---- CARBOU (Rangifer tararzdus) 957. u! \.eew.aun :o. ('. 63pr /"" \""t Rc,-~ ~! ~:itis. C. A '..: and thennj J4 ~-+5~85t'..:oiogica anj h. id anima, '! '!ie~. i Landor., ab. Buiere /omadism. \\'.,pp. Chemica imtammas. A h and Wid. t povement patsh and Game, J e of anters in it. Deer Soc. /rhou in nonhf niw condri utting of rein 'nim. Beha\. i tons affecting j.enzie i'ons. Dept. of i Br.. Ottawa. :>en!being an ne Sanctuary. md an outine )ept. of nter Jttawa. 53pp. ichen species. t uibou. Pages Kein. and R. and Caribou Rep. no.. >se with M99 24:94-0. of skin in the ysio. 7:355- Monographic pzig. 264pp Sea)ona on of capnve , "to surviva in >ia. Northwest t the fubarctic u herd. Am. uibou_studies. ), 2. 48pp. --- C,6?.. Tie migratory barren-ground caribou of Canada. Can. Wid!. Serv. Monogr. no. 3. Ottawa. 34pp nteraction between hurren-ground caribou and muskrats. Can. J. Zoo!. 48: Warbe fy distribution among some Canadian caribou. Pa2e~ J. R. Luick. P. C. Lent. D. R. Kein. a~d R. G. White. eds. Proc. st nt. Reindeer and Caribou Symp. Bio. Pap. Univ. Aaska. Spec. Rep. no.. 55pp. Kein, D. R Range-reated differences in growth of deer refected in skeeta ratios. J. Mamma. 45C2): a. Food seection by North American deer and their response to overutiization of preferred pant species. P~~es in A. Watson. ed. Anima popuations in reation to their food resources. Br. Eco. Soc. Symp. no. 0. Backwe Sci. Pub., Oxford and Edinburgh. 447pp b. nteractions of Rangifer tarandus (reindeer and caribou) with its habitat in Aaska. 8th nt. Congr. Game Bio., Hesinki, Finand. Pp Kein, D. R., and White, R. G Parameters of caribou popuation ecoogy in Aaska. Proc. Symp. and Workshop, Bio. Pap. Univ. Aaska. Spec. Rep. no pp. Krog, J. 0.. and Wika. M The circuation ~n the -growing reindeer anters. Pages in J. R. Luick, P. C. Lent. D. R. Kein, and R. G. White, eds. Proc. st nt. Reindeer and Caribou Symp. Bio. Pap. Univ. Aaska. Spec. Rep. no.. 55 pp. Kuropat, P Range interreationships of the western arctic herd. Aaska Coop. Wid. Res. Unit, Univ. Aaska. Semiannua Prog. Rep. 30: Kuyt. E Food habits of woves on barren-ground caribou range. Can. Wid!. Serv. Rep. Ser. no pp. LaPerriere, A. J., and Lent. P. C.. Caribou feeding sites in reation to snow characteristics in northeastern Aaska. Arctic 30: Lent, P. C. 965a. Observations on anter shedding by femae barren~ ground caribou. Can. J. Zooi. 43: b. Rutting behavior in a barren-ground caribou popuation. Anim. Behav. 3: a. Caving and reated socia behavior in the barren-ground caribou. Z. Tierpsycho. 23: b. The caribou of northwestern Aaska. Pages 48!-57 in N. J. Wiimovsky and J. N. Wofe. eds. Environment of the Cape Thompson region, Aaska. U.S. AtQmic Energy Comm., Washington, D.C.,250pp Mother-infant reationships in unguates. Pages 4-55 in V. Geist and F. Wathers, eds. The behaviour of unguates and its reation to management. VoL. UCN New Ser. Pub!. no. 24. Morges, Switzerand. 940pp A review of acoustic communication in Rangifer rarandus. Pages in J. R. Luick, P. C. Lent, D. R. Kein, and R. G. White, eds. Proc. st nt. Reindeer and Caribou Symp. Bio. Pap. Univ. Aaska, Spec. Rep,. 55pp, Lentz, C. f',, and Hart, J. S The effect of wind and moisture on heat. oss through the fur of newborn caribou. Can. J. Zoo!. 38: Leopod, A, S,, and Daring, F. F. 953, Widife in Aaska. Ronad Press Co. New York. 29pp, Lewin, V,, and Stefox, J, G Functiona anatomy of the tai and associated behavior in woodand caribou. Can. Fied Nat. : Lon!!. F. M.. ed.. First nternationa Conference on Widife Biotcemctrv. Pa!!cs -5'> m Proc. nt. Conf. Wid!. Bioteemetry: Lar;mie. Wyoming Juy Second mernationa Confer:nce on Widife Brotcemetrv. Page~ -259 in Pro~. nt. Conf. Wih.. Bioteemet~.. L.;amie. Wvomin!! Juv. and August JQ79. Loomis. F. G Dentition of aniodacty!.. Bu. Geo. Soc. Am. 36: Luick. J. R. and White. R. G Gucose metaboism m femae reindeer. Pages in J. R. Luick, P. C. Lent. D. R. Kein, and R. G. White, eds. Proc. st nt. Reindeer and Caribou Symposium. Bio. Pap. Univ. of Aaska. Spec. Rep.. 55pp. McEwan. E. H Seasona annui in the cementum of the teeth of barren-ground caribou. Can. J. Zoo!. 4: Hematoogica studies of barren-ground caribou. Can. J. Zoo. 46: Twinning in caribou. J. Mamma. 52:479. McEwan. E. H., and Whitehead, P. E Seasona changes in the energy and nitrogen intake in reindeer and caribou. Can. J. Zoo!. 48: Reproduction in fetnae reindeer and caribou. Can. J. Zoo!. 50: McEwan. E. H., and Wood, A. J Growth and deveopment of the barren-ground caribou. Pan : Heart girth. hind foot ength. and body weight reationships. Can. J. Zoo!. 44:40-4. McEwan. E. H.: Wood. A. J.: and Nordan. H. C Body temperature of barren-ground caribou. Can. J. Zoo. 43: McGowen, T. A Caribou studies in northwestern Aaska. Proc. Western Assoc. State Game and Fish Commissioners 46: Manning, T. H The reationship of the Peary and barren-ground caribou. Arctic nst. North Am. Tech. Pap pp. Mier. D. R Seasona changes in the feeding behavior <f barren-ground caribou on the Tatga winter range. Pages in V. Geist and F. Waters. eds. The behaviour of unguates and its reation to management. Vo. 2. UCN New Ser. Pub. no. 24. Morges, Switzerand. 940pp Bioogy of the Kaminuriak popuation of barr~n-ground caribou. Part 3: Taiga winter range reationships and diet. Can. Wid. Serv. Rep. Ser. no pp. Mier, F. L mmobiization of free-ranging backtaied deer with succinychoine choride. J. Wi<!. Manage. 32: Eruption and attrition of mandibuar teeth in barren-ground caribou. J. Wid!. Manage. 36: a. Age determination of caribou by annuations in denta cementum. J. Wid!. Manage. 38:47~ b. A new era: are migratory barren-ground caribou and petroeum expoitation compatibe? Trans. Northeastern Sec. Wid!. Soc. 3: c. Bioogy of the Kaminuriak popuation of barren-ground caribou. Part 2: Dentition as an indicator of sex and age: composition and sociaization of the Popuation. Can. Wid. Serv. Rep. Ser. no pp Numbers and distribution of Peary caribou on the arctic isands of Canada. Pages 6-9 in D. R. Kein and R. G. White, eds. Parameters of caribou popuation i

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